Journal of the Botanical Research Institute of Texas.

Botanical
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Journal of the Botanical Research Institute of Texas

J. Bot. Res. Inst. Texas ISSN 1934-5259

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Table of Contents

SYSTEMATICS

MISSOURI BOTANICAL

DEC 0 9 2010

Supplemental notes on Bolivian Xyris (Xyridaceae) GARDEN LIBRARY

Robert Kral

Emmet J. Judziewicz, Eddie L. Shea, and Tanya M. Wayda
A new Leucophyllum (Scrophulariaceae) from Sonora, Mexico
James Henrickson and Thomas R. Van Devender

Poa ramifer (Poaceae; Pooideae: Poeae; Poinae), a new aerially branching gynomonoecious
species from Peru

Robert J. Soreng and Paul M. Peterson

Two new Andean species of Solanum section Crinitum (Solanaceae)

Frank T. Farruggia, Michael H. Nee, and Lynn Bohs

A new species of Campomanesia (Myrtaceae) from Bahia, Brazil, based on specimens
collected by J.S. Blanchet over 150 years ago
Leslie R. Landrum and Marla Ibrahim U. de Oliveira

Folia taxonomica 18. The status of Passiflora citrifolia and a new species in subgenus
Astrophea (Passifloraceae), Passiflora jussieui
Christian Feuillet

Folia taxonomica 19. Typifications in Dilkea (Passifloraceae)
Christian Feuillet

Taxonomic notes on the genera Stenotis
Hedyotis greenei to Stenotis

Edward E. Terrell and Harold Robinson

sfer of

Transfer of Hedyotis intricata to Arcytophyllum (Rubiaceae)

Edward E. Terrell and Harold Robinson

Nomenclatural transfers in the genus Myrsine (Myrsinaceae) for New Caledonia
Jon M. Ricketson and John J. Pipoly III

Hedychium forrestii (Zingiberaceae) with a new synonymy and a variety from India
E. Sanoj, M. Sabu, and T. Rajesh Kumar

The reinstatement of Ptilimnium texense (Apiaceae) and a new key to the genus

A new variety of Bromus flexuosus (Poaceae; Pooideae; Bromeae; sect. Bromopsis)

Ana Maria

Planchuelo

The genus Rytidosperma (Poaceae) in tl
Stephen J. Darbyshire, Henry E. Connor, ^

: United States of Arne
ID Barbara Ertter

Typifications of names in Agalinis, Gerardia, and Toman
J.M. Canne-Hilliker AND John F. Hays

A new combination in Lolium perenne (Poaceae; Poeae); L. pei
Joseph K. Wipff, III

anchaceae)

Paleocharis nearctica gen. and sp. nov (Cyperaceae) in Cretaceous Canadian amber
George O. Poinar, Jr. and David J. Rosen

563

569

581

587

595

603

609

615

625

627

633

653

663

677

683

685

Coleataenia Griseb. (1879): the correct name for Sorengia Zuloaga & Morrone (2010)
(Poaceae: Paniceae)

Robert]. Soreng

Congruence between allometric coefficients and phylogeny in stipoid grasses; an evo-devo

study

Jack Maze

Thomas Walter’s species of Hedysarum (Leguminosae)

Daniel B. Ward

FLORISTICS, ECOLOGY, AND CONSERVATION

Listado floristico y aspectos ecoldgicos de la familia Poaceae para Chihuahua, Durango y
Zacatecas, Mexico

Yolanda Herrera Arrieta y Armando CoRTfis Ortiz

Mirandea grisea (Acanthaceae), new for Coahuila and Durango, Mexico
Eduardo Estrada-CastillOn, Jos£ Angel Villarreal-Quintanilla, and Jorge Arturo Alba-Avila
Registro de dos nuevas localidades y reubicacion de individuos de una de ellas de Astrophytum
myriostigma (Cactaceae) en Durango, Mexico

Jaime SAnchez Salas, Gisela Muro P^rez, Eduardo Estrada CastillOn y Mario GarcIa Aranda
Erigeron mancus (Asteraceae) density as a baseline to detect future climate change in La
Sal Mountain habitats
James F. Fowler and Barb Smith

Scallopleaf sage (Salvia vascyi: Lamiaceae) discovered in Arizona

James W. Cain, 111, Brian D. Jansen, Richard S. Felger, and Paul R. Krausman

Paspalum pubiflorum and P. quadrifarium (Poaceae) new to California, with a key and

notes on invasive species

Richard E. Riefner, Jr., Silvia S. Denham, andJ. Travis Columbus
Sarracenia minor var. okefenokeensis (Sarraceniaceae) discovered outside of the
Okefenokee Swamp area
Jacob S. Thompson

Polycarpon tetraphyllum (Caryophyllaceae) new to the flora of Louisiana
Charles M. Allen, Jarrod Grandon, Krisztian Megyeri, and Brad Waguespack
Additions to the vascular flora of New Mexico
Ben S. Legler

693

705

711

739

741

747

755

761

771

775

777

Book Reviews and Notices 568, 580, 586, 608, 618, 624, 640, 652, 682, 746, 754, 774, 776, 785

Reviewers for Volume 4 (2010)— 787
Index to Volume 4 (2010)

Titles of Articles with Authors— 788
Authors— 791

Botanical Names and Subjects — ^792

Pipoly,

Myrsine oblanceolata (M. Schmid) Ricketson &
Pipoly, comb, nov.— 630

Myrsine oblanceolata subsp. doensis (M. Schmid)
Ricketson & Pipoly, comb, et stat. nov. — 631
Myrsine obovalifolia (M. Schmid) Ricketson &
Pipoly, comb. nov. — 631

Myrsine ouameniensis (M. Schmid) Ricketson &
Pipoly, comb, nov — 631

Myrsine ouazangonensis (M. Schmid) Ricketson &
Myrsine ovicarpa (M. Schmid) Ricketson & Pipoly,

Ricketson & Pipoly, comb, et sta
Myrsine parvicarpa subsp. pachypbylla (M. Schmid)
Ricketson & Pipoly, comb, et stat. nov. — 631
Myrsine poumensis (M. Schmid) Ricketson & Pipoly,
comb. nov. — 631

Myrsine pronyensis (Guillaumin) Ricketson &
Pipoly, comb. nov. — 631

Myrsine spissifolia (M. Schmid) Ricketson & Pipoly,

Myrsine tcbingouensis (M. Schmid) Ricketson &
Pipoly, comb. nov. — 631

Myrsine verrucosa (M. Schmid) Ricketson & Pipoly,

Myrsine verrucosa subsp. microphyUa (M. Schmid)
Ricketson & Pipoly, comb, et stat. nov. — 631
Myrsine yateensis (M. Schmid) Ricketson & Pipoly,
comb. nov. — 632

Paleocharis Poinar & D.J. Rosen, gen. nov. — 686

Passiflora jussieui Feuillet, sp. nov — 61 1
Poa ramifer Soreng & PM. Peterson, sp. nov. — 587
. Nee&Farruggia, sp. nov. —

ee Farruggia, sp.

nov — 596

Stenotis greenei (A. Gray) Terrell & H. Rob., comb.

SUPPLEMENTAL NOTES ON BOLIVIAN XYRIS (XYRIDACEAE)

Robert Krai

Herbarium (VDB)

Botanical Research Institute of Texas
500 E 4th Street

Fort Worth, Texas 76102-4025, U.SA.

ABSTRACT

RESUMEN

tinct and are not found in existing treatments (Krai 1988, 1994; Maguire & Smith 1964; Smith & Downs
1968). These are presented below.

1. Xyris crassifunda Krai, sp. nov. (Fig. 1). Type: BOLIVIA. Santa Cruz. Velasco Prov.; Parque National Noel Kempff M. Los

Plants slender, rushlike, caespitose, hard-based, perennial, (5-)6-7(-7.6) dm high. Roots fibrous. Shoots
short, stout, hard, close-set, arising from a ligneous, caudex-like base, stem bases often hidden by persistent,
dark fibrils from old leaf bases. Principal leaves erect or ascending, flexuous and twisted, 35-45 cm long,
longer than the scape sheaths; sheaths entire or basally ciliate, Vs-Vi of blade length, the convex, several-
carinate base of outer (lower) principal leaves, more dilated, often ciliate, more abruptly narrowed distally
than those of the upper, all at very base castaneous or dark red-brown, smooth, upsheath becoming paler,
transversely rugose or rugulose medially, the smooth, chartaceous borders gradually narrowed, apically
converging to an acute ligule 3-3.5 mm long; blades narrowly linear, twisted and flexuous, 1-1.5 mm thick
at level of ligule, gradually narrowed distally to a subulate-conic tip, coarsely and unevenly few-costate,
costae making rounded angles, these and the uneven shallow sulcae yellow-green or tan, transversely finely
papillose-rugulose. Scape sheaths firm, rounded-costate, yellow-green, papillose, conduplicate, smooth and
red-brown proximally, open apically, the chartaceous edges converging as ligule, the apex projecting just
beyond as a stubby, conic, apiculate blade. Scapes erect, twisted, proximally 2-2.5 mm thick and lustrous
red-brown, distally narrowing to ca. 1.5 mm thick, coarsely low-costate, increasingly transversely rugu-
lose-papillate, dull yellow-green. Mature spikes several-flowered, obovoid, 1-1.2 cm; bracts firm, spirally

564

hair (right). L. Capsule, one valve removed. M. Seed.

565

imbricate, backs convex, ecarinate, dark red-brown with distinct, paler, narrowly elliptic dorsal area ca. 2
mm long; sterile bracts few, ovate to broadly obovate, transitional to fertile, the lower two ca. 5 mm, ovate,
these obovate to oblong, 5. 5-7.5 mm, the distal narrowest, all apically rounded, borders narrow, scarious,
entire, becoming shallowly erose. Lateral sepals free, inequilateral, narrowly elliptic to oblanceolate, the
firm keel proximally ciliolate or entire, medially increasingly crisped ciliate, apically crisped-fimbriate.
Petal blades elliptic-obovate, ca.7 mm long, irregularly low-toothed apically, broadly acute, yellow. Anthers
lance-linear, ca. 3 mm, apically shallowly emarginate, base sagittate, on filaments ca. 2 mm; staminodia
bibrachiate, 3 mm, including dense brushes of moniliform beard hairs. Capsule narrowly oblong-ellipsoid,
ca. 5 mm, placentation basal. Seeds irregularly ellipsoid, ca. 1 mm, dull red-brown with darker coarse.

anastomosing, irregular ribs.

Distribution . — Known only from a meseta slope, on a sandy clay substrate by gallery fore;

Bolivia. Represented solely by the type collection.

Material of this xyrid shows traits both of Xyris rigida Kunth, Enum. Pi. 4:15, 1843, and ^

Malme, Bih. Kongl. Svenska Vetensk.-Akad. Handl. 24, Afd. 3, No. 3: 10, plate 1, fig. 7, 1898, both of th
planaltan Campos of southern Brazil, thus part of that complex of tall, slender, rush-like perennials whos
leaves have distinct auricles and whose spike bracts produce distinct dorsal areas. Unfortunately it share

b uave uibLiiK.1 aulicicb aiiu wiiubc spike bracts produce distinct dorsal areas. Unfortunately it shares
of the distinguishing characters of both, but the characters are combined differently. Since t
only example gotten thus far, it may be that intermediates will turn up as eastern Bolivia is further <
but until such happens it seems best to introduce this as a species.

Etymology.— The name “crassifunda” refers to the thickened caudex-like base.

Arroyo, Mostacedo, Guillen, Gutierrez, Zapata y Pena s.n. (holotype; USZ;

Stout-based, caespitose perennial 5-7(-8 est.) dm, the stems contracted, hard. Leaves erect or in narrow
fans, the outermost mostly scale-like, bladeless. Principal leaves 4-5 cm, sheaths ca. Vz-Vi as long as blades,
entire, abruptly dilated, convex, multicostate at base, castaneous, distally keeled, gradually narrowed,
eligulate, to blade; blades linear, flattened, 2-3 mm wide, shallowly several-nerved, surface smooth with
lines of elliptic punctae, margins narrowly incrassate, densely scaberulous, narrowed to a narrowly acute,
somewhat thickened, apex. Scape sheathes tubular, lustrous red-brown, twisted, short-bladed, shorter
than leaves. Scapes erect to ascending, twisted, proximally ca. 1.5 mm, distally ca. 1 mm thick, rounded
to oval in cross-section with low, irregular ribs, smooth save for lines of narrowly elliptic punctae and one
strong, scaberulous costa. Spikes obovoid, ca. 1.2 cm long, of many, spirally imbricate, convex, brownish or
reddish-brown, lustrous bracts without evident dorsal areas, base attenuate, of many sterile bracts, the lower
ones smallest, broadly ovate, ca. 2 mm, grading upward to larger, progressively longer, more oblong fertile
ones; fertile bracts prevalently oblong-ovate to oblong, 5.5-7 mm, those of midspike and above narrowest in
outline, apically rounded-emarginate, erose or entire, and with a prominent, though narrow, raised midvein
cresting a shallow apical angle. Lateral sepals free, inequilateral, oblong-lanceolate to oblong-oblanceolate.

Journal of the Botanical Research Institute of Texas 4(2)

566

567

ca. 7 mm long, the narrow, firm, shallowly sigmoid-curved keel mostly antrorsely or retrorsely pilosulous-
ciliate medially, antrorsely ciliolate distally. Petal blades (est.) oblong-elliptic, ca. 5 mm, bluntly acute,
yellow. Anthers lance-oblong, 2-2.5 mm, emarginate and sagittate, on stout filaments ca. 2 mm. Capsules
lanceoloid, ca. 5 mm, light brown; placentation basal. Seeds asymmetrically ellipsoid, 0. 8-0.9 mm, dark
red-brown, not farinose, longitudinally finely multiribbed.

Distribution . — Known thus far only from the type locality, one of humid grasslands around forested

islands and amongst termit
In character of habit,
spikes are longer, attenuate
character. Many of the mid
midnerves (apically often d;

ite mounds.

leaf, and scape this example fits Xyris metallica Klotsch ex Seub. However, the
: (thus narrower-based), and with several more sterile bracts of distinctly different
i and upper bracts of a spike have distinct, if shallow, keels apically, with strong
iarker-pigmented) marking the crests of that shallow angle. The lateral sepals and

Etymology.— The name “submetallica” is applied here against the possibility that connecting morpholo-
gies will prove a relationship.

ACKNOWLEDGMENTS

The kindness of curators as well as that shown by the original collectors is very gratefully acknowledged.
Richard Carter (VSC) kindly assisted with technical aspects of preparing the manuscript. I kindly thank
two anonymous reviewers for their helpful comments.

REFERENCES

Kral, R. 1 988. The genus Xyr/s (Xyridaceae) in Venezuela and contiguous northern South America. Ann. Missouri
Bot. Card. 75:522-722.

Kral, R. 1994. Xyridaceae. In: Flora of the Guianas, Ser. A: Phanerogams, fasc. 15. Koeltz Scientific Books. Konig-
stein, Germany.

Maguire, B. and LB. Smith. 1964. Xyridaceae. In: B. Maguire, J.J. Wurdack, and collaborators. The botany of the
Guayana Highland— Part V. Mem. New York Bot. Gard. 10(5):7-37; figs. 1-22.

Smith, L.B. and R.J. Downs. 1968. Xyridaceae. In: F.C. Hoehne, FI. Brasilica 9(2):1-215.

568

Journal of the Botanical Research Institute of Texas 4(2)

BOOK REVIEW

WoifGANG Stuppy AND RoB Kesseler. 2008. Fruit: Edible, Inedible, Incredible. (ISBN 978-1-554-07405-1.
hbk). Firefly Books Inc., P.O. Box 1338, Ellicott Station, Buffalo, New York 14205, U.S.A. (Orders:
service@fireflybooks.com, 1-800-387-5085, 1-800-565-6034 fax). $60.00, 264 pp., selection of color
images, preface, and appendices, 11 x 12

J. Hot Res. Inst Teas 4(2); 568. 2010

TWO NEW BOLIVIAN SPECIES OF AULONEMIA
(POACEAE; BAMBUSOIDEAE: BAMBUSEAE)
Emmet J. Judziewicz Eddie L Shea

Robert W. Freckmann Herbarium
Department of Biology and Museum of Natural History
University of Wisconsin-Stevens Point
Stevens Point, Wisconsin 54481, U.SA

Department of Biology
University of Wisconsin-Su
Stevens Point, Wisconsin 54481, USA.
eshea241@uwsp.edu

Tanya M. Wayda

Department of Biology
University of Wisconsin-Stevens Point
Stevens Point, Wisconsin 54481, U.S.A.
tanya.m.wayda@uwsp.edu

ABSTRACT

RESUMEN

INTRODUCTION

Aulonemia Goudot (Poaceae: Bambusoideae: Bambuseae: Arthostylidiinae) is a genus of about 34 species of
woody bamboos (McClure 1973; Calderon and Soderstrom 1980; Clayton and Renvoize 1986; Judziewicz
et al. 1999; Judziewicz et al. 2000). Of the 34 species that we currently recognize within the genus, 20 have
been described since 1990 (Clark 2004; Clark et. al. 2007; Clark and Londono 1990; Clark et al. 1997;
Judziewicz 2005; Judziewicz et al. 1991; Judziewicz and Riina 2005; Judziewicz and Tyrrell 2007; Renvoize
1998). The Andean countries of Venezuela, Colombia, Peru, and Bolivia appear to have many new species
of Aulonemia awaiting description Qudziewicz, pers. obs.).

Recent collections from the Parque Nacional Madidi (http://www.mobot.org/MOBOT/Research/madidi/)
on the northern slope of the Andes in the Department of La Paz, in western Bolivia have revealed a number
of distinctive new species of Aulonemia, two of which are described below. Specimens were examined from

570

several herbaria (F, ISC, K, MO, P, US, UWSP and WIS), but only MO had representatives of the new taxa.
Plant parts were measured using a mm ruler, and the abaxial leaf blade epidermis of both species was ex-
amined using dried material in a Hitachi S3400 scanning electron microscope. Neither taxon is included
in the most recent treatment of Bolivian grasses (Renvoize 1998; 38-41). The following key differentiates

IS with internodes al

It or poorly-developed, if present then lacking
A. scripts Judz., Shea & Wayda

A. boliviana Renvoize

5. Branches of inflorescence i

nemia bromoides Judz. & Shea, sp. nov. (Figs. 1, 2, 3D-F). Type: BOLIVIA. U Paz: Prov. Franz Tamayo: Sen
Apolo-San Jos^ de Uchupiamonas, a media hora de 3 de Mayo, hacia Tumia, 14->33.654'S [-39.2-], [=04.6"], 18

Miranda 6- Freddy Qmqui 289 (holotyTe: LPB!; isotypes: MO 4793775! 47937761).

n longae, glabra, si

Apparently cespitose from sympodia
men occupying 40-70% of the diameter of the culm; culms apparently dimorphic, some with well-developed
foliage leaves and terminal inflorescences, others with poorly-developed culm leaves, no foliage leaves, and
terminating in inflorescences, the culms thus apparently dimorphic. Culm leaves with sheaths 3-18 cm
long, 0.8-1.2 cm wide (folded width), glabrous, with a minute inner ligule and no outer ligule or auricles,
efimbriate, bladeless or with tiny erect blades ca. 2.5 mm long and 1.5 mm wide. Culms with lower internodes
glabrous, slightly purplish and verrucose, the upper internodes slightly striate, glabrous or with scattered
glassy appressed hairs 0.7-1. 5 mm long. Branching pattern not seen. Foliage leaves in complements of at
least 3, efimbriate; sheaths slightly striate, glabrous or with scattered glassy appressed hairs 0.7-1. 5 mm
long, stramineous with tiny gray or purple maculae, lacking an auricle at the summit; outer ligules abaxi-
ally 1 mm long, laterally 1.5-2 mm long; inner ligules 20-25 mm long, foliaceous, linear-lanceolate, pale
stramineous (at least when mature); pseudopetioles 4-6 mm long; blades 18-25 cm long, 5.3-6.7 cm wide,
lanceolate-ovate, obtuse to slightly cuneate at the base, acuminate at the apex, reflexed, glabrous, the margins
cartilaginous and somewhat antrorsely hispid. Inflorescence an ovoid panicle 25-35 cm tall and 17-22 cm
wide with the base included in the uppermost leaf sheath; rachis and primary branches smooth, glabrous,
and slightly maculate, the primary branches ascending at a 45° angle, the secondary branches and pedicels
capillary and antrorsely scabrous, the pedicels up to 6 cm long. Spikelets 40-43 mm long, 3.0-4. 1 mm wide,
robust, stramineous with prominent grayish-green nerves, glabrous, the bracts all acute and awnless with
pubescent margins; lower glume 2. 7-4.4 mm long, 0.8-1.3 mm wide (folded), lanceolate to lance-ovate,
3-5-nerved, some of the lateral nerves not extending to the apex; upper glume 4.8-6.7 mm long, 1.0-1.6
mm wide (folded), lanceolate, 5-7-nerved, some of the lateral nerves not extending to the apex; lowermost
floret sterile, lacking a palea, 6.1-10.1 mm long, 1.1-2.0 mm wide (folded), lanceolate, 7-9-nerved; second

572

574

near base, the margins densely fringed with erect clear cilia ca. 0.5 mm long; stamens 3, the filaments ap-
parently dilated, the anthers 4.5 mm long, 0.8 mm wide, linear, stramineous to brown. Fruit 4.3-5.9 mm
long, 0.7-1.0 mm wide, narrowly oblong, rusty brown; embryo 2.9 mm long, 0.8 mm wide; hilum 4.1-5 .4
mm long; all fruits observed were immature.

Leaf anatomy of abaxial leaf blade surface (Fig. 3A-C; terminology follows Ellis 1979).

Costal zones . — Spaced ca. 150 pm apart.

575

Fk. 4. Aulonemia scripta [Maldonado et al. 3 100, MO). Detail of specimen showing newsprint adhering to and imprinted upon culm and leaves.

Papillae. — In general very abundant; up to 12 pm long and 6 pm wide, variable, occasionally slightly
branched at the apex.

Stomates. — Common; shape unknown, possibly triangular, nearly obscured by overarching papillae;
each stomate overarched by 9-15 papillae, each papillae 4-12 pm long; stomates in 2 rows flanking each
costal zone and slightly extending into the intercostal zone.

Interstomatal cells. — Shape apparently rectangular, often nearly obscured by papillae; papillae absent
or solitary, globose to slightly oblique.

Long cells. — Rectangular, 40-60 pm long, 4-6 times as long as wide; outline of cells sinuous; papillae
simple, in one or two rows, globose.

Prickles. — ^Abundant, 10-20 prickles/mm in 1-2 rows in both costal and intercostal zones, individual
prickles 40-110 pm long, base at least twice as long as the stomates, the barb shorter than the base.

Short cells. — Not evident, perhaps obscured by over-arching papillae.

Microhairs. — Common; two-celled, the basal cell 60-80 pm long, 10-12 pm wide, the apical not seen,
apparently detached in SEM preparation.

Macrohairs. — None seen.

Known only from the type locality in Parque Nacional Madidi, La Paz, Bolivia, Aulonemia scripta is
distinctive in its apparently viscid hollow culms, viscid leaves, and awned spikelets. The specific epithet of
the new species alludes to the newsprint that is imprinted upon and adhering in fragments to the culm and
leaves of the type collection. The culms, branches, leaf sheaths, and lower leaf blade surfaces also have a
shiny “varnished” appearance suggestive of viscidity. The collectors did not note any stickiness on their label
notes, but this is not surprising: In another new species of Aulonemia from Brazil (Pedro Viana, pers. comm,
from field observations) with viscid foliage (and spikelets), the collector, the late Thomas R. Soderstrom,
who took copious field notes on the bamboos he collected, does not note any viscidity, and viscidity is not
apparent in his dried specimens Qudziewicz, pers. obs.). Besides Aulonemia scripta and the unpublished

A

f. C. Lower glume. 0. Upper glume. E. Lemma.

578

new Brazilian species alluded to above, only two other described members of the Bambusoideae are known
to have viscid foliage. Both are congeners and Costa Rican endemics (Pohl 1980; Pohl and Davidse 1994).
Aulonemia viscosa (A. Hitchcock) McClure also has viscid foliage, but it also has viscid inflorescences (includ-
ing spikelets), is more robust in stature (4-5 m tall and 5-15 cm in diameter), has pulpy, succulent, easily
compressed culms, efimbriate leaves with broader blades, and purple -maculate awnless spikelets. In dried
herbarium specimens (e.g., Pohl & Lucas 13113, F), the only evidence of viscidity is the shiny, varnished
appearance of the summit of the leaf sheath.

Aulonemia patriae R.W. Pohl, also from Costa Rica, differs from A. scripta in its larger spikelets, leaves
with well-developed fimbriae 15-30 mm long, and ovate leaf blades; in a dried herbarium specimen (Pohl
12798, F), there is no evidence of viscidity. It is not clear whether viscid foliage arose once or several times
independently in all of these species; the morphological variation among them is quite diverse. The only
other report of viscidity in a member of the Bambusoideae is in Chusquea spencei Ernst, a high elevation
species of the p^iramos of Colombia and Venezuela, in which the culm internodes are frequently sticky
(Clark 1989: 58). In other plants, viscidity has been postulated to be an attribute of what might be termed
“protocarnivory” (Darwin 1875; Spomer 1999; Chase et al. 2009).

The three described “viscid” species of Aulonemia can be distinguished by the following key:

1 . Culms ca. 0.5 m tall, 2-7 mm in diameter, rigid and hollow; spikelets with lemma awns 4-5.5(-7) mm long;
foliage leaves narrowly lanceolate, 2-2.7 cm wide; leaf fimbriae sparse {0-2 per leaf), delicate, 4-8 mm long;

1 . Culms 4-5 m tall, 5-1 5 mm in diameter, easily crushed, pulpy and succulent; spikelets awnless; foliage leaves
narrowly to broadly ovate, 4-8 cm wide; leaf fimbriae either absent or well-developed (15-30 mm long);

2. Leaves with fimbriae well-developed, 1 5-30 mm long, the blades broadly ovate, 4-8 cm wide; peduncles

and inflorescence branches not viscid; spikelets not maculate, 25-40 mm long A. patriae

2. Leaves efimbriate, the blades narrowly ovate, 4-5 cm wide; peduncles and inflorescence branches viscid;

spikelets purple-maculate, 15-25 mm long A. viscosa

Aulonemia scripta is assessed as Critically Endangered because its EOO (extent of occupancy) is (as far as is
lUCN (2001) applies.

ACKNOWLEDGMENTS

We thank UWSP student Eva C. Hathaway for the line drawings, Sol Sepsenwol for assistance in scanning
electron microscopy, Gerrit Davidse and James Solomon for the loan of specimens from MO, Virginia Freire
for assistance with the Spanish abstract, and Lynn G. Clark and Ximena Londono for helpful reviews, and
Monica Moraes (Directora, Herbario Nacional de Bolivia) for confirming the LPB holotypes.

REFERENCES

Calderon, C.E. andT.R, Soderstrom, 1 980. T

and comments. Smithsonian Contr. Bot. 44:1 -27.

Chase, M.W., MJ.M. Christenhusz, D. Saunders, and M.F. Fay. 2009. Murderous plants: Victorian Gothic, Darwin and
modern insights into vegetable carnivory. Bot. J. Linnaean Soc. 1 61 :329-356.

Clark, LG. 1989. Systematics of Chusquea section Swallenochloa, section Verticillatae, section Serpentes, and
section Longifoliae (Poaceae: Bambusoideae). Syst. Bot. Monogr. 27:1-1 27.

Clark, LG. 2004. New species of Aulonemia and Chusquea (Poaceae: Bambusoideae: Bambuseae) from south-
eastern Brazil. Rev. Bras. Bot. 27:31-36.

Clark, LG., EJ. JuDziEwta, and C.D. Tyrrell 2007. Aulonemia ximeniae (Poaceae: Bambusoideae: Bambuseae), a new
species from Colombia and Venezuela. Bamboo Sci. Cult. 20:1-6.

Clark, LG. and X. Londoi^o. 1990. Three new Andean species of Aulonemia (Poaceae: Bambusoideae). Ann. Mis-
souri Bot. Card. 77:353-358.

580

BOOK NOTICE

Lincoln Taiz and Eduardo Zeiger. 2010. Plant Physiology, 5th Edition. (ISBN 978-0-878-93866-7, hbk.).
Sinauer Associates, Inc., 23 Plumtree Road, Sunderland, Massachusetts 01375, U.S.A. (Orders: www.
sinauer.com). $124.95, 782 pp., 593 illus., selection of color figures, SEM images, graphs, tables, im-
ages and appendices, 9" x 11 14".

lation regarding plant physiology. The authors include a variety
lany processes that occur within plants. Taiz and Zeiger added

tion included on the companion website (www.plantphys.net).

A NEW LEUCOPHYLLUM (SCROPHULARIACEAE) FROM SONORA, MEXICO

James Henrickson

Thomas R.Van Devender

Plant Resources Center
University of Texas
Austin. Texas 7871 2, U.5A

Sky Island Alliance
738 5th Avenue
Tucson. Arizona 85705, U.S.A.

ABSTRACT

INTRODUCTION

from near Alamos, Sonora, Mexico has revealed a distinctive new species of Leucophyllum. It
other Leucophyllum species in all critical features, e.g., in having exstipulate leaves, dendritic
stiture, ebracteate flowers with separate sepals, didynamous stamens, partially adnate filaments,
1 parallel anther sacs with the outermost sac connected across the anther tip, and sympetalous,
weakly zygomorhic, blue-purple corollas that are orange spotted on the throat floor and pilose

All other species of Leucophyllum occur in habitats marginal to and above deserts, (in mixed desert
scrub, yucca woodland, izotal, thorn scrub) from trans-Pecos, central and southeastern Texas southward
through east and central Mexico to Oaxaca (Henrickson & Flyr 1985). However, the new taxon occurs far
to the west in a seasonally more mesic, usually frost-free oak woodland on the west slopes of the Sierra
Madre Occidental in southeastern Sonora (Martin et al. 1998). And while all other species of Leucophyllum
are strongly branched, sometimes thorny shrubs with small leaves, the new species is sparsely, more openly
branched, with thicker young stems, larger leaves, flowers and fruit, all features showing adaptation to this

DESCRIPTION

LeucophyUum mojinense Henrickson &r T.R. Van Devender, sp. nov. (Fi^. 1, 2). Type: MEXICO. Sonora: l

above; young stems ± 2 mm diameter, terete, white pannose with dense, appressed, thin-walled, branched
hairs; older stems 3-5 mm diameter, with vestiture tardily diminished; basal stems to 1.4 cm diameter,
periderm gray to dark gray; internodes 2-9 mm long. Leaves alternate, subopposite, rarely some opposite,
erect-ascending, thickish, elliptical, broadly elliptical to obovate, ovate in sucker shoots, (25-)32-56 mm
long, ll-22(-38) mm wide, acute to obtuse-rounded, abruptly apiculate at the tips, narrowly cuneate above

J. Bot Res. Inst Texas 4(2): 581 - 585. 2010

Journal of the Botanical Research Institute of Texas 4(2)

solitary, axillary; pedicels 2-5 mm long, 1-1.2 mm wide, pannose; calyx 9-11 mm long; sepals 5, lanceolate,

keeled, tapered to thickish, obtuse to acute tips, ± pannose-floccose outside, stipitate glandular inside, the
inner sepals with shaggy matted marginal hairs to 1 mm long, the sepals straight and persisting around
fruit. Corollas ascending, 35-40 mm long, the basal tube (that enclosed in the calyx) 6-7 mm long, ± 4-5
mm diameter, thickened, narrowly invaginated where adnate to filament bases, expanding above sepals to
a broadly, slightly ampliate throat 18-22 mm long, to 8-11 mm in diameter, the 5 lobes rounded, 9-12 mm
long, 9-10 mm wide, entire, rounded or the abaxial most minutely apiculate at the tip, spreading-upcurved
(2 adaxial, 2 lateral and one abaxial), glabrous except where pilose with unicellular hairs to 2 mm long, 0.1
mm wide on the abaxial lobe and distal-mid tube-throat within, the tube whitish, striped with blue-purple
outside, with rows of orange spots on the mid-basal, abaxial corolla throat within, the lobes stronger blue-
purple; stamens 4, included, didynamous, (the adaxial-most, fifth stamen absent); adaxial (posterior-lateral)
paired filaments 18-21 mm long, adnate to corolla tube base for 10-11.5 mm, the free filaments 8-10 mm

long ± 1.3 X 0.7 mm thick, positioning the introrse anther along the root ot the corolla throat about o-c
mm below the corolla lobes, the abaxial (anterior-lower) filaments 15-17 mm long, adnate to corolla tube
base for 8-10 mm, the free filament 6-8 mm, ± 0.5 mm in diameter, positioning the introrse anthers at the
base floor of the tube throat proximal to the other pair, the filaments white, glabrous; anthers cream white
anther sacs oblong, the adaxial pair 4.5-5 mm long, ± 1.2 mm in total width, the abaxial pair ± 2.5 mn
long, 1-1.2 mm in total width, the anther sacs initially parallel, the outer sac continuous around the tip, the
paired anther sacs diverging at anthesis, those of the adaxial stamens drying peltately explanate. Ovarie:
superior, weakly stipitate glandular near the tip; styles 19-21 mm long, glabrous, exserted well beyond the
anthers; stigmas narrowly compressed lanceolate, 1.5-2.8 mm long, to 0.7 mm wide, acute, opening a
anthesis, stigmatic along the margins. Fruit brownish, ligneous, broadly ovoid, ± 8 mm long, to 5-6 mn
wide, acuminate, opening along 4 distal sutures; seeds numerous, yellowish-tan, irregularly angular b
compression, blunt at the ends, 0.9-1 mm long, 0.4-0.5 mm wide, the surfaces with a fine hexogonal pat
tern. All flower measurements are from rehydrated herbarium material.

(data from Google Earth, 2010).

DISCUSSION

Leucophyllum mojinense differs from other species of Leucophyllum in many features. Vegetatively it has a
more open branching with prominent stems and large leaves (see the stems in the background of Fig. lA).
The stem and leaf vestiture consists of narrow-based, asymmetrically inflated cylindrical cells with very

(Fig. 2). In leaves the upper cells are horizontally suspended above broad, erect basal cells. The flowers are
the largest in the genus ranging from 35-40 mm in overall length, with an extended, open throat. Like
other species in the genus the stamens are didynamous, basally epipetalous, with the larger adaxial anthers
positioned at the top of the corolla throat, opening downward, and the smaller abaxial anthers positioned
at the base of the corolla throat, opening upward. But unlike other species the thickish basal corolla tube is
deeply invaginated opposite the attachment of the filament bases. Also, the adaxial anthers are the largest
in the genus and the style tip is narrowly lance-ovate.

Within Leucophyllum, the new species shares most characteristics with L. hintonorum Nesom, a species
with cuneate-based leaves from gypsum habitats between Aramberri and Zaragoza, Nuevo Lebn. But while
L. hintonorum has moderately large leaves 20-35 mm long, and moderately large flowers 16-28 mm long
with orange spots in the lower throat, it differs strongly in vestiture, growth habit, and habitat making any
statement of true relationships tentative.

Etymology.— The specific epithet refers to the southern Sonora term for tropical deciduous forest, monte
mojino, the dominant vegetation visible in the Rio Cuchujaqui drainage below the type locality. Mojino refers
to the reddish-grey color of the dry season forest. It also celebrates the name of the land where it grows.
Rancho Ecological Monte Mojino (REMM) and the efforts of Nature and Culture International, Inc. to pre-
serve this threatened habitat.

The new species was discovered by Martin Gabriel Figueroa-Martinez on July 15, 2005 as part of a
vegetation survey of the Rancho Santa Barbara area at a site about 21 km and 66° east-northeast of Alamos
in southern Sonora (Martin et al. 1998). This area is on the 6,000 hectare REMM owned by Nature and
Culture International, Inc., a private non-profit organization based in San Diego, California. REMM was
established in 2004 to protect tropical deciduous forest within the large (92,890 hectares) Area Protegida
de Fauna and Flora Sierra de Alamos-Rio Cuchujaqui.

Rancho Santa Barbara is in a very interesting area on the western edge of the Sierra Madre Occidental.
It is in the Rio Cuchujaqui drainage, a major tributary of the Rio Fuerte in Sinaloa. Tropical deciduous for-
est occurs along most of the elevational gradient in REMM from 350 m along the Cuchujaqui to 1500 m
above Santa Barbara. The type locality of L. mojinense, at 1200 m elevation, is in oak woodland just above
tropical deciduous forest and below pine-oak forest. Two large populations are in same area on rocky poor
soils referred to as sabanias. Associated species include Quercus spp., Dodonaea viscosa, Agave bovicornuta,
Lysiloma watsonii, and Ipomoea arborescens. Dario Sauceda, who made the first collections of the taxon, noted
that he has not seen the plant elsewhere in the Sierra Madre Occidental in this area in Sonora or in western
Chihuahua (pers. comm. 2009 to Stephanie A. Meyer). The type of the new Erigeron barbarensis Nesom & T.R.
Van Devender was from Arroyo Santa Barbara 1.5 km west-northwest of the L. mojinense populations (Nesom
& Van Devender 2007) and type locality of Hesperaloe tenuifolia G.D. Starr lies 1.7 km. south-southwest of
the type locality of L. mojinense on Cerro Agujudo (Starr 1997).

ACKNOWLEDGMENTS

We thank Stephanie A. Meyer, Martin Gabriel Figueroa-Martinez, and Dario Sauceda of Rancho Ecolbgico
Monte Mojino for calling this new species to our attention and providing specimens. We thank Mark A
Dimmitt and Ana Lilia Reina- Guerrero for their comments and encouragement and Paul Fryxell for the
Latin translation, Fernando Chiang for Spanish translation, Kanchi Ghandi for nomenclatural advice,
Robert Harms for aligning the figures, and Richard S. Felger and an anonymous reviewer for comments on

REFERENCES

Henrickson, J. and D. Fiva 1985. Systematics of Leucophyllum and Eremogeton (Scrophulariaceae). Sida 11:
107-172.

Martin, P.S., D.A. Yetman, M. Fishbein, P. Jenkins, T.R. Van Devender and R.K. Wilson (eds.). 1 998. Gentry's Rfo Mayo plants.

TheTropical Deciduous Forest & Environs of Northwest Mexico. University of Arizona Press, Tucson.

Nesom, G.L 1991. A new species of Leucophyllum (Scrophulariaceae) from Nuevo Leon, Mexico. Phytologia
71:337-339.

Nesom, G.L. andTR. Van Devender 2007. A new species of Erigeron (Asteraceae: Astereae) from the Rfo Mayo Region
of Sonora, Mexico. Phytologia 89:21 9-222.

Starr G.D. 1 997. A revision of the genus Hesperaloe (Agavaceae). Madrono 44:282-296.

Hill ill

586

BOOK REVIEW

Tracy Disabato-Aust. 2008. 50 High-Impact, Low-Care Garden Plants: Tough-but-Beautiful Plants
Anyone Can Grow. (ISBN 978-0-881-92950-8, pbk.). Timber Press Inc., The Haseltine Building, 133
S.W Second Avenue, Suite 450, Portland, Oregon 97204-3527, U.S.A. (Orders: orders® timberpress.
com, 1-800-827-5622). $16.95, 168 pp., 135 color photos, 7 V2" x 9".

POA RAMIFER (POACEAE: POOIDEAE; POEAE: POINAE),

A NEW AERIALLY BRANCHING GYNOMONOECIOUS SPECIES FROM PERU

Robert J. Soreng and Paul M. Peterson

National Museum of Natural History

Washington, DC 20013-7012, U.S.A.
sorengr@si.edu; peterson@si.edu

ABSTRACT

On a high ridge above a tributary north of the canyon of the Rio Santa along the western slopes of the Andes
in the Provincia de Corongo, Departamento Ancash of northern Peru, the authors in 2008 collected a Poa
that was undoubtedly a new species. The type collection was gathered about eight km above Pilipampa,
on a winding road between Bambas and Miraflores, below the pass between the provinces of Corongo and
Pallasca. About 100 plants were noted in the one population we found on the south facing shrubby slopes
between 2750 and 3040 m in elevation. No other stations are known for the species, but we expect the

animals or otherwise minimally disturbed. We have examined collections of Poa from South America,
particularly from Peru at CPUN, MO, MOL, US, and USM [acronyms follow Thiers (2009)], but found no
other collections matching the new species.

Poa ramifer Soreng & PM. Peterson, sp. nov. (Fi^. 1 A-C, 2 A-O). Type: PERU. Departamento Ancash: Provincia

Plants perennial, cespitose from broad based clumps (to 1 m diam.) with extravaginal primary shoots from
a knotty base; primary roots 0.4 mm diam.; primary culms 80-100 cm tall (including panicles), erect or
slightly decumbent, 1-1.5 mm diam. at base, smooth, perenniating with 7-10-leaf bearing nodes, nodes
smooth, glabrous, with intravaginal and extravaginal leafy secondary shoots arising from nodes 5-60 cm
above base along the primary flowering culms, secondary shoots flowering in the next season and developing
a few new leafy tertiary shoots. Leaves mostly cauline; sheaths slender, slightly compressed, scabridulous,
papery (not becoming fibrous), lower sheaths closed up to the ligule, upper sheaths 2-4 times as long as
their blades, closed ca. 40% of their length, lightly scabridulous; blades 2-15 cm long, 1-2 mm wide, flat
or folded on drying, thin, shallowly keeled, abaxially and adaxially scabridulous mostly over the costae,
with 6 or 7 narrow and well spaced costae (intercostal area 1-3 times as wide as each costal rib) on either

Journal of the Botanical Research Institute of Texas 4(2)

Fk. 1. ftwnimfftf Soreng & P.M. Peterson [PM Peterson & RJ. Soreng 21804 (US)]. A. Habit. B. Lateral extravaginal shoot with cataphyll. C Panicle.

^ species of Poa from Peru

589

590

side of the midrib, adaxially with single rows of bulliform cells flanking the midrib; ligules 2-4 mm long,
membranous, whitish, usually with a few deep lacerations, abaxially scabrous, those of the lower shoots
connate in front, of the upper shoots grading to long decurrent, apex acute. Panicles 4-10 cm long, erect
except for nodding tip, open, sparse with 20-65 spikelets, lowest internodes 1.8-2.6 cm long; branches 1 or
2(-3) per node, capillary, 0.1-0.2 mm in diam., flexuous, widely spreading to steeply reflexed, sparingly to
moderately scabridulous along the angles, longest 2. 5-5. 5 cm long with 3-10 spikelets in the distal halves;
pedicels shorter than the spikelets, scabridulous. Spikelets (2-)3-4(-5)-flowered, laterally compressed,
proximal (l-)2 or 3 florets perfect, distal 1 or 2 florets pistillate; glumes 2.2-3. 2 mm long, unequal, keeled,
smooth or lightly scabridulous, edges smooth to closely scabridulous, apex acute; lower glumes 2. 2-2.4
mm long, lanceolate, 1-nerved; upper glumes 2.9-3.2 mm long, 0.9-1.35 mm wide, ca. twice as wide as
lower, broadly lanceolate 3(or 4)-nerved; rachilla internode between first and second floret ca. 0.6 mm long,
densely scabrous, distal internodes ca. 1.1 mm long, moderately to densely scabrous, terete; callus laterally
compressed in the dorsal side, smooth, glabrous or infrequently with a few dorsal hairs, the hairs to 0.8
mm long; lemmas 3. 3-3. 8 mm long, 0.7-1. 1 mm wide, laterally compressed, strongly keeled, membranous,
5-nerved, marginal veins moderately pronounced, intermediate veins obscure to moderately pronounced,
margins involute and narrowly white scarious, keel and marginal veins densely scabridulous, keel glabrous
or sericeous with 1-several hairs near the base, the hairs to 0.4 mm long, surface between veins lightly to
moderately scabridulous, silica cells common, surface between the marginal vein and edge moderate to
densely scabridulous, edge smooth or sparsely to moderately scabridulous, apex acute; paleas equaling
lemma in length, keels densely scabridulous, between keels moderate to densely scabridulous, flanges
smooth, about as wide as the gap between the keels; lodicules 2, obtuse, glabrous, in perfect florets lodicules
0.2-0.3 mm long, unlobed or briefly lobed, in pistillate florets lodicules 0. 5-0.6 mm long, lanceolate with
a distinct lateral lanceolate lobe; stamens 3; anthers 2.0-2.8 mm long, or vestigial, then ca. 0.2 mm long;
ovary glabrous, styles 2, terminal, adjacent, stigmas ca. 1.5 mm long, lanceoloid, white, with moderately to
densely plumose branches from the base, primary branches sparingly rebranched; caryopses 1.7-2 mm long,
narrowly lanceolate, hard, greenish, tightly adherent to the palea, hilum ca. 0.2 mm long, narrowly elliptic.

Distribution and Habitat.— Shrubby upper slopes of the canyon of the Rio Santa around Cerro Santo
Toribio, with Eragrostis lurida]. Presl, E. pilgeri Fedde, E. weberbaueri Pilg., Festuca sp., Melica scabra Kunth,
Salvia sp., Baccharis sp., and other Asteraceae shrubs, 2750-3040 m.

Comments— Poa ramifer might be closely related to the more widespread P. aequatoriensis Hack, that
occurs in relatively low elevations (for Poa) in Colombia, Ecuador, and Peru. In Peru the latter species occurs
on the Amazonian side of the Andes as far south as Ayacucho in more mesic western slopes from 2300-3500
m (Tovar 1993). Poa aequatoriensis is a more mesophilic species that occasionally stools, roots at the nodes,
and occasionally branches along the lower culms, but it does not branch well up into the old erect culms as
in P. ramifer (Figs. 1 <Sr 2). In addition, P. aequatoriensis has panicles with ascending branches, smaller spike-
lets with narrower glumes (up to 0.5 mm wide), lemmas (up to 0.7 mm wide) that are distinctly pubescent
along the keel, and calluses that are distinctly webbed (contrast with Figs. 1 & 2). Both species apparently
exhibit the same type of gynomonoecism (type Ib, see discussion below), as is common in northern Andean
species of Poa, but the fertile anthers in P. aequatoriensis are only about 1 mm long.

DISCUSSION

The unusual branching habit within Poa ramifer (Figs. lA & B; 2A) could be the result of natural selection
favoring longevity of culms in the lower elevation habitats in Peru where few species of Poa occur. Of the
2990 grass collections PMP with collaborators have made in Peru between the years 1997 and 2008, 421
collections are of Poa (not including Aphanelytrum, Dissanthelium, Tovarochloa; see Gillespie et al. 2008: Al-
though these genera are resolved within Poa in DNA analyses, they have not yet been transferred). Of these
only eight Poa collections (including P. ramifer) were made below 3000 m, and of these only three were
found below 2900 m. Even though species of subfamily Chloridoideae often dominate the lower elevations

of the coastal slopes of the Andes, vegetation is extremely sparse below 2000 m, except in riparian habitats.
These open, low elevation slopes are evidently extremely xeric and not well suited for Poa. Temperatures at
the type locality of Poa ramifer probably never drop below freezing [location of the type is well within the
low of 0-4°C minimum temperature zone, and near the 4-8°C minimum; Servicio Nacional de Meteorologia
e hidrologia del Peru (2009) website map], and cloud banks regularly occur in this zone (we saw a cloud
bank and photographed it during our trip) providing humidity, dew, and infrequent rains, at least during
the wet season. This habitat may have favored the development of above ground culms that persist in this

1953, e.g., Oxalis gigantea Barneoud).

There are now 34 species of Poa (including P. ramifer) accepted for Peru by Soreng et al. (2003, updated
on-line version, see 2009b). We predict that the new species belongs to the large, world-wide Poa subg. Poa
supersect. Homalopoa (Dumort.) Soreng & L.J. Gillespie based on its absence of rhizomes, relatively closed and
compressed sheaths, distinct lemma nerves, and scabrous palea keels (preliminary nuclear rDNA sequence
data supports this placement; Lynn Gillespie, pers. comm.). Based on morphological characteristics, all native
Peruvian species of Poa belong to supersect. Homalopoa. DNA sequences of 2/3 of the indigenous Peruvian
Poa species (not including the new species) have been phylogenetically examined (Refulio-Rodriguez 2007;
Gillespie et al. 2008; L. Giussani, L. Gillespie, M.A. Negritto, N.F. Refulio-Rodriguez, R.J. Soreng unpubl.
data), and none have aligned outside of the group that we interpret as P. supersect. Homalopoa. The new
species is most obviously distinct from all other species of Poa of the Americas in its peculiar branching
habit. Individual plants of P. ramifer form large broad-based clumps (to 1 m diam at base), with 7-10-noded
erect culms reaching one meter in height that are partly supported by shrubs (Fig. 3). A panicle terminates
the main culm or terminates lateral shoots that originate well up along the previous season’s culm (unusual
within the genus). A similar aerial branching habit is known from a few Australian species of Poa supersect.
Homalopoa sect. Brizoides Pilg. s.l. (e.g., P. homomalla Nees), and a few others in P. sect. Brizoides subsection
Australopoa Soreng, L.J. Gillespie & S.W.L. Jacobs (e.g., P. tenera Hook, f., P. Jugicola D.I. Morris, and P. or-
thoclada N.G. Walsh; Gillespie et al. 2009, Soreng et al. 2009a), but this habit has presumably been derived
independently in Peru. Unlike two other South American species of Poa (P. hieronymi Hack, and P. myriantha
Hack.) with long straggling multi-noded culms to 2 m, P. ramifer has anthers that are 2 mm long in the
perfect florets (versus stamens 1 mm long in the perfect florets), calluses that are glabrous (infrequently with

1 or 2 short hairs present; versus web well developed), panicles that are 5-10 cm long with 20-65 spikelets
(versus 20-36 cm long panicles with more than 100 spikelets), and erect culms that perenniate with new
flowering culms arising from the upper nodes.

The assessment of the breeding system of the new species is a bit preliminary. Although the species was
more common upslope, we only collected specimens from a few plants of P. ramifer as these were not com-
mon where we stopped. The plants collected were gynomonoecious, within the spikelets with the proximal

2 or 3 florets perfect and the distal 1 or 2 florets pistillate. Apparently the distal pistillate flowers produce
seed, but only partially developed caryopses were found in the “perfect” flowers. This arrangement of sexes
appears to fit one of four types of gynomonoecism detailed by Connor (1979; type Ib as outlined by Soreng
& Keil 2003) where spikelets consistently contain proximal perfect florets and distal pistillate florets. This
type of gynomonoecism is present in at least 20 South American species of Poa (Anton & Connor 1995;
Negritto & Anton 2000), although a few South American species of the genus exhibit a separate type called
sequentially adjusted (as opposed to fixed types recognized by Connor 1979) gynomonoecism (Soreng &
Keil 2003; e.g. P.fibrifera Pilg., P. plicata Hack., P. horridula Pilg.), wherein the frequency of pistillate flow-
ers and spikelets increases through the season in some proportion of individuals within a population (best

)rth American species such as P. cuspidata Nutt, and P. tracyi Vasey). Further study of
viduals of P. ramifer is needed to confirm whether the species is fixed or sequentially gynomonoecious
eng & Keil 2003), as there is extensive variation in breeding systems in Poa (Anton & Connor 1995),
1 within gynomonoecious species (Connor 1979; Negritto & Anton 2000; Soreng & Keil 2003).

592

Journal of the Botanical Research Institute of Texas 4(2)

Fit 3. Habit of Poa ramifer [PM Peterson & RJ. Soreng2im (US)] in situ. Photo by R J. Soreng.

593

This is the first time in Poa, so far as we are aware, that different lodicule forms within a spikelet have
been noted between florets. In P. ramifer the lodicules are present but reduced in the proximal perfect florets
in each spikelet, while these structures are fully developed in the distal floretfs) within the same spikelet. In
the perfect florets the lodicules are short, blunt and unlobed (Fig. 2] & M), whereas lodicules of the pistil-
late florets are lanceolate and lobed (Fig. 2L) as is typical of most species of Poa. We know of no previous
documentation of morphological dimorphism of the lodicules between pistillate and perfect florets within
the same species of Poa. In the Chilean dioecious species, P. cumingii (sect. Dioicopoa E. Desv.) Trin., lodicules
are absent or poorly developed in staminate florets and well developed in pistillate florets (RJS, pers. obs.).
Lodicules in monoecious and dioecious species in some grass genera (e.g., Phams and Leptaspis, and Boute-
loua, respectively) are well developed in staminate florets but absent in pistillate florets Oudziewicz 1987;
Kinney et al. 2008). In some gynodioecious species of Cortaderia, lodicules are longer in pistillate plants
than in hermaphroditic plants (Connor 1973). These patterns seem to be diverse. Lodicules are thought to
be involved in opening the grass floret at flowering. Why lodicules should be morphologically dimorphic
between proximal perfect and distal pistillate florets within the same spikelet in this new species of Poa,
and whether the pattern in P. ramifer occurs in related species, requires further study.

The habit of the new species is quite the opposite of the newly described Poa unispiculata Davidse, Soreng
& P.M. Peterson (2010). The latter species grows in the Altiplano of Departamento Pasco, Peru, at ca. 4400

single spikelet and upwards of 200 inflorescences per individual. Poa unispculata is gynodioecious with some
plants that are pistillate and other plants that have perfect spikelets. As different as these two species may
appear (see Davidse et al. 2010, Fig. 3), they have some similarities in growth form. The dwarf species (P.
unispiculata) has many culm leaves/nodes (averaging 20-30) along the flowering culm with extremely short
internodes (ca. 1 mm long). In this species the initial culms are prostrate or ascending, emerging within

of the older nodes. Just above some of the older nodes a succession of additional intravaginal leafy shoots
are produced and many of these form new inflorescences, and so on, as the mat expands over time. Do the
similarities in branching habit indicate a close phylogenetic relationship between these two Peruvian spe-
cies? We hope to test this hypothesis by adding P. ramifer and P. unispiculata to our growing library of DNA
sequence for the genus.

ACKNOWLEDGMENTS

We thank the National Geographic Society Committee for Research and Exploration (grant number 8087-06)
for field and laboratory support; the Smithsonian Institution’s, Restricted Endowments Fund, the Scholarly
Studies Program, Research Opportunities, Atherton Seidell Foundation, and Biodiversity Surveys and Invento-
ries Program, all for financial support; Alice R. Tangerini for illustrating the new species and pointing out the
dimorphic lodicules!; Christian Feuillet for help preparing the Latin diagnosis; Lynn Gillespie for preliminary
DNA ITS and ETS sequence analyses; Zulma Rtigolo de Agrasar and Joseph Wipff for reviewing the manuscript;
and the following colleagues who facilitated or assisted with fieldwork: Asuncion Cano Echevarria, Maria
I. LaTorre, Nancy Refulio-Rodriguez, Jenny Rojas Fox, Isidoro Sanchez Vega, and Dorita Susanibar Cruz.

REFERENCES

Anton, A.M. and H.E. Connor 1995. Floral biology and reproduction in Poa (Poeae: Gramineae). Austral. J. Bot.
43:577-599.

Connor H.E. 1973. Breeding systems in Cortaderia (Gramineae) Evolution 27:663-678.

Connor H.E. 1979. Breeding systems in the grasses: a survey. New Zealand J. Bot. 17: 547-574.

Davidse, G., RJ. Soreng, and P.M. Peterson. 2010. Poa unispiculata, a new gynodioecious species of cushion grass
from Peru with a single spikelet per inflorescence (Poaceae, Pooideae, Poeae, Poinae). J. Bot. Res. Inst. Texas.
4:37-44.

'I Peru. Estac. Exp. Agricola "La

Ferreyra, R. 1953. Comunidades de vegetales de algunas lomas c
Molina,” Bol. 53:1-88.

Gillespie, LJ., RJ. Soreng, R.D. Bull, S.W.L. Jacobs, and N.F. REFuuo-RoDRfGUEZ. 2008. Phylogenetic relationships in su btribe
Poinae (Poaceae, Poeae) based on nuclear ITS and plastid trnT-trnb-trnF sequences. Botany 86:938-967.
Gillespie, LJ., RJ. Soreng, and S.W.L. Jacobs. 2009. Phylogenetic relationships of Australian Poa (Poaceae: Poinae),
including molecular evidence for two new genera, Saxipoa and Sylvipoa. Austral. Syst. Bot. 22:41 3-436.
JuDziEwicz, EJ. 1 987. Taxonomy and morphology of the tribe Phareae. PhD Dissertation, University of Wisconsin,

Madison.

Kinney, M.S., J.T. Columbus, and E.A. Fariar 2008. Unisexual flower, spikelet, and inflorescence development in mo-
noecious/dioecious Bouteloua dimorpha (Poaceae, Chloridoideae). Amer. J. Bot. 95:1 23-1 32. 2008

Negritto, M.A. and A.M. Anton. 2000. Revision de las especies de Poa (Poaceae) del noroeste argentine. Kurtziana
28:95-136.

Refulio-RodrIguez, N.F. 2007. Systematics of DissantheliumJnn. PhD Dissertation, Claremont Graduate University,
Claremont, California.

Servicio Nacional de METEOROLOGfA E HiDROLOGfA DEL PerO (SENAMFII). 2009. httpy/www.senamhi.gob.pe/ (web-site
maps accessed Dec. 2009).

Soreng, RJ., LJ. Gillespie, and S.W.L. Jacobs. 2009a. Saxipoa and Sylvipoa - two new genera and a new classification
for Australian Poa (Poaceae: Poinae). Austral. Syst. Bot. 22:401 -41 2.

Soreng, RJ., LM. Giussani, and M. Negritto. 2003 [updated on line]. Poa L. In: RJ. Soreng, P.M. Peterson, G. Davidse,
EJ. Judziewicz, F.O. Zuloaga,T.S. Filgueiras, and O. Morrone, eds. Catalogue of New World grasses (Poaceae):
IV. subfamily Pooideae. Contr. U.S. Natl. Herb. 48:505-580.

Soreng, RJ. and D. Keil 2003. Sequentially adjusted sex-ratios in gynomonoecism, and Poa diaboli (Poaceae), a
new species from California. Madrono 50:300-306.

Soreng, RJ., G. Davidse, P.M. Peterson, F.O. Zuloaga, EJ. Judziewicz, T.S. Filgueiras, and 0. Morrone. 2009b. Catalogue of
New World grasses. httpy/www.tropicos.org/ProJect/CNWG [2 Oa 2009, continually updated]

Thiers, B. 2009. Index Herbariorum: A global directory of public herbaria and associated staff. New York Botanical
Garden's Virtual Herbarium. httpY/sweetgum.nybg.org/ih/ [continually updated]

Tovar, O. 1 993. Las Gramineas (Poaceae) del Peru. Ruizia 1 3:1 -480.

ANDEAN SPECIES OF SOLANUM SECTION CRINITUM (SOLANACEAE)

Michael H. Nee

596 Journal of the Botanical Research Institute of Texas 4(2)

are restricted to tropical South America, with highest diversity along the wet tropical eastern slopes of the
Andes.

Solanum cyathophorum M. Nee & Farruggia, sp. nov. (Fig. 1). Type: ECUADOR. Napo: 8 km no abajo de Puerto
Misahualli, por el Rio Napo y 1.5 km al sur, 01“04'S. 77“36'W. 450 m. 25 May 1985, D. Neill, W. Palacios &J. Zaruma 6506
(holotype; QCNE; isotypes: MO, NY, QAME).

Tree 3-10(-20) m x ca. 60 cm dbh. Trunk with sharp, stout broad-based prickles, the bark light tan to dark,
wrinkled in vertical ridges, the slash soft, white with yellow fibers; flowering stems unarmed, glabrous to
densely pubescent with sessile to short-stalked light tan multangulate-stellate hairs, the multangulate apex
0. 1-0.3 mm in diameter, the rays 7-10+. Sympodial units difoliate, geminate. Leaves simple, the blades 15-30
X 7-15 cm or more, ca. 2 times as long as wide, elliptic to lanceolate, cbartaceous to subcoriaceous, slightly
discolorous, the fresh and dried leaves dark green and somewhat shiny adaxially, lighter green to golden-tan
abaxially, the adaxial surface nearly glabrous to moderately pubescent with stalked reduced-stellate hairs,
the stalks ca. 0.1-0.3 mm, multiseriate at the base, rays l(-2), eglandular, the abaxial surface mostly densely
pubescent with golden-tan sessile to multiseriate-stalked porrect-stellate hairs, the stalks 0-0.03 mm, the
rays 5-8, the midpoints absent; major veins 8-9 on either side of midvein; base cuneate to oblique; margin
entire to shallowly repand; apex acute to acuminate; petioles (2-)4-5 cm, glabrous to densely pubescent
with hairs like those of the young stems. Inflorescences 2.5-4.5 cm, extraaxillary, unbranched or branched,
with 8-15 flowers, the plants strongly andromonoecious, with one to few hermaphroditic flower(s) at the
base of the inflorescence and all other flowers functionally staminate, the axes densely stellate-pubescent
with hairs like those of the stems, unarmed; peduncle 2-5 mm; rachis 2. 3-4.2 cm; pedicels 10-15 mm in
flower and fruit, densely congested, spaced 1-4 mm apart, articulated at base. Flowers 5-merous. Calyx ca.
4.5 mm long, the tube at anthesis 1. 1-1.4 mm, the lobes ca. 2.5 x 2 mm, the apex truncate to broadly obtuse,
the abaxial surface densely pubescent with short-stalked to sessile porrect-stellate hairs, unarmed; fruiting
calyx tube becoming slightly inflated and knobby, the lobes 5-5.5 x 3-3.9 mm, slightly reflexed, subtending
the fruit. Corolla 4.5-5 cm in diameter, 22-23 mm long, stellate to rotate-stellate, lobed for more than half
of its length, membranaceous, violet to blue, the tube 6-7.5 mm, the lobes 11-14 x 3-3.5 mm, lanceolate,
with moderate interpetalar tissue, sparsely pubescent adaxially with sessile porrect-stellate hairs, the rays
5-8, the midpoints often pronounced, ca. 0.1 mm long, densely pubescent abaxially along central portion
of lobes with sessile porrect-stellate hairs. Stamens equal, the filament tube 0-0.1 mm, the free part of the
filaments 1.5-1.8 mm, glabrous; anthers ca. 13 x 2.8 mm, tapered, not connivent, yellow, the pores directed
distally, not opening into longitudinal slits with age. Ovary glabrous to sparsely pubescent with stalked
glandular hairs; style in hermaphroditic flowers 14-15 x 0.2-0. 5 mm, cylindrical, emergent from anther
column, curved at apex, glabrous or sparsely pubescent in lower half with sessile stellate or short-stalked
unbranched glandular hairs; style in staminate flowers 4-4.5 x ca. 0.2 mm, cylindrical, included within
anther column, straight at apex, glabrous or sparsely pubescent in lower half with sessile stellate or short-
stalked unbranched glandular hairs; stigma capitate, slightly bilobed. Fruit a berry, 1-1.9 cm in diameter,
globose, apparently green and juicy at maturity, glabrous, the pericarp thin. Seeds 2.6-3 x ca. 2.5 mm,
strongly flattened, reniform, orange to light brown, rugose.

Distribution and phenology.— Clearings and open places in disturbed, transitional and lowland tropical

into southern Colombia and northern Peru. Flowering specimens were collected in February-August and
October-December. Fruiting specimens were collected in February, May, August- September and November-
December.

Conservation status . — According to the lUCN Red List Categories (lUCN 2010), S. cyathophorum is clas-

598

Journal of the Botanical Research Institute of Texas 4(2)

sifted as LC (Least Concern). Populations of this species are known from multiple locations throughout a
broad range, and many of the collections are from protected areas.

Local names.— Ecuador: apumpu (Quichua, Hurtado 2733); apumpu yura (Neill et al. 6506); papa mandi
(Freire et al 2175); untukar (Shuar, Shiki RBAE222); bamba (Mowbray 702102); ttottopaje (Cofan, Ceron

Uses.—Ceron 301 reports that the bark is used to induce vomiting.

This is the species listed as “sp. nov. ined.” in Nee (1999) under unnamed series 1 of Solanum section
Crinitum. Within section Crinitum, S. cyathophorum most closely resembles S. altissimum Benitez and S.
kioniotrichum Bitter ex J. F. Macbr. Solanum altissimum is distributed throughout the western Amazon Basin
from Colombia and Venezuela to Peru and western Brazil. Solanum kioniotrichum is endemic to Departmento
Loreto, Peru. All three species have small round glabrous fruits (1-2 cm) and predominantly entire leaves.
Solanum cyathophorum differs from S. altissimum and S. kioniotrichum in having a calyx with truncate lobes
that do not completely cover the corolla in bud, thus exposing the end of the corolla. In S. altissimum and
5. kioniotrichum the calyx lobes are acute to obtuse and completely cover the corolla in bud, only splitting
later to expose the corolla. The adaxial leaf surfaces of S. cyathophorum are either completely glabrous or
sparsely pubescent with hairs having a multiseriate stalk and 1-2 horizontal apical ray cells. Specimens

reduced to rounded knobs. Solanum kioniotrichum can be easily distinguished from both 5. altissimum and
S. cyathophorum by the pronounced basal cells of the adaxial leaf hairs. Evidence from nrDNA ITS sequence
data and morphology suggests that S. cyathophorum is most closely related to 5. altissimum (F.T. Farruggia
& L. Bohs, unpublished data).

Etymology— The name refers to the shape of the calyx: cyatho- (from the Greek “kyathodes”), meaning
cuplike, and the suffix -phorum, meaning to bear or carry.

del Guamues y Puerto Asis, 270 m. 21 Dec 1940, J. Cuatrecasas 11245 (COL, F, US). ECUADOR. Morona-Santiago: Gualaquiza Cantdn.

Solanum adenobasis M. Nee & Farruggia, sp. nov. (Fig. 2). Type: PERU. Dept. Amazonas: Bagua, near Puente Almendro
1. km 296 of Marafton road (from Olmos junction), 5“15'S. 78°20'W, 620 m, 6 Jul 1984, S.D. Knapp &J. Mallet 6565 (holotype:

599

Tree 3-10(-20) m x ca. 6-12 cm dbh. Trunk with sharp, stout broad-based prickles, the bark light tan to
dark brown, thin, the slash soft, white with yellow fibers; juvenile stems densely covered with stout broad-
based prickles; flowering stems unarmed or with broad-based prickles, often densely pubescent with long
multiseriate-stalked porrect-stellate hairs, the stalks 4-5 mm long, the base of the stalk with a tuft of small
glandular hairs, the stellate apex 2-3 mm in diameter, the rays 3-6, the midpoint ca. 0.5 mm long. Sympodial
units difoliate, geminate. Leaves simple, the blades 30-120 x 15-45 cm or more, ca. 2 times as long as wide,
ovate to lanceolate, chartaceous, the fresh leaves dark green and somewhat shiny adaxially, lighter green
to golden-tan abaxially, the adaxial surface moderately pubescent with multiseriate-stalked porrect-stellate
hairs, the stalks 1-2.5 mm, the stellate apex 1-2 mm in diameter, the rays 2-4(-6), eglandular, the abaxial
surface densely pubescent with golden-tan multiseriate-stalked porrect-stellate hairs, the stalks ca. 1 mm, the
rays 5-6, the midpoint lacking; major veins 5-6 on either side of midvein; base cuneate to oblique; margin
shallowly lobed to nearly entire, the lobes deltoid; apex acute to acuminate; petioles 6.5-9(-30) cm, moder-
ately to densely pubescent with hairs like those of the young stems. Inflorescences 2.5-8 cm, extraaxillary,
unbranched or once-branched, with 8-15 flowers, the plants strongly andromonoecious, with one to few
hermaphoditic flower(s) in the inflorescence and all other flowers functionally staminate, the axes densely
stellate-pubescent with hairs like those of the stems, unarmed; peduncle 0.5-3 cm; rachis 2-7.5 cm; pedicels
5-17 mm in flower and fruit, densely congested, spaced 1-4 mm apart, articulated at base. Flowers 5-mer-
ous. Calyx 12-15 mm long, the tube at anthesis 1.1-2 mm, the lobes 11-14 x 2.5-3 mm, the apex acute to
acuminate and thick, the abaxial surface densely pubescent with long-stalked porrect-stellate hairs, the stalks
2-3 mm with glandular hairs present at base, unarmed; fruiting calyx tube becoming strongly inflated and
knobby, the lobes 10-35 x 8-18 mm, subtending the fruit. Corolla 4-6(-8) cm in diameter, rotate-stellate,
lobed for more than half its length, membranaceous, violet fading to white, the tube 20-23(-28) mm, the
lobes 29-36(-39) x 6-7 mm, debate, with abundant interpetalar tissue, the adaxial surface glabrous or with
sparse sessile to short-stalked stellate to multangulate hairs along midvein near apex, the abaxial surface
densely covered along lobes with sessile porrect-stellate hairs. Stamens equal, the filament tube 0-0.1 mm,
the free part of the filaments 1.7-2 mm, glabrous; the anthers 11-14 x 1. 8-2.1 mm, tapered, connivent,
yellow, the pores directed distally. Ovary glabrous; style in hermaphoditic flowers 12-15 x 0.5-0. 8 mm,
cylindrical, emergent from anther column, curved at apex, sparsely pubescent at base with short glandular
hairs; style in staminate flowers 2-3.2 x ca. 0.2 mm, cylindrical, included within anther column, straight
at apex, sparsely pubescent at base with short glandular hairs; stigma short-cylindrical to capitate. Fruit a
berry, 4-7 cm in diameter, globose, green and juicy at maturity, glabrous and lustrous, the pericarp thin.
Seeds 2.5-3.5 x 2.5-3 mm, strongly flattened, reniform, orange to light brown, rugose.

Distribution and phenology.— Clearings and open places in disturbed, lowland to upland tropical rain-
forest, 180-1 200C-1600) m in elevation, mainly along the eastern Andean slopes in southern Ecuador and
northern Peru. Flowering specimens were collected March-December. Fruiting specimens were collected
May-December.

Conservation status . — According to the lUCN Red List Categories (lUCN 2010), S. adenobasis is classified
as VU-Bla+biii; A2c; D1 (Vulnerable). Populations of this species are located near expanding population
centers leading to highly fragmented populations. The extent of occupancy is estimated to be less than
25,000 km^ and there are estimated to be less than 1,000 mature individuals across its range. There is also
a continuing decline in suitable habitat in these regions due to deforestation and the establishment of new
settlements.

Local names. — Ecuador: pungala (Narvaez 446); untukar (Shuar, Kunkumas RBAE184). Peru: untukag
(Ancuash 460).

Uses . — Used as a graft with the edible-fruited naranjilla, Solanum quitoense Lam. (Narvaez 446). Accord-
ing to Kunkumas RBAE184 the fruit is not edible.

X. Drawn from photos of L Bohsetal. 3808 8,3833 (UT).

oG. D'Arcyrthe legacy of a

Journal of the Botanical Research Institute of Texas 4(2)

lUCN Standards and Petitions Subcommittee. 2010. Guidelines for using the lUCN Red List categories and criteria.
Version 8.0. Prepared by the Standards and Petitions Subcommittee in March 2010. Downloadable from
http://intranet.iucn.org/webfiles/doc/SSC/RedList/RedListGuidelines.pdf.

Levin, R.A., N.R. Myers, and L. Bohs. 2006. Phylogenetic relationships among the "spiny solanums" {Solanum sub-
genus Leptostemonum). Amer. J. Bot. 93:157-169.

Linnaeus, C. 1753. Species plantarum. Stockholm.

Nee, M. 1999. Synopsis of Solanum in the New World. In: Nee, M., D.E. Symon, R.N. Lester and J.P. Jessop, eds.

Solanaceae IV: Advances in biology and utilization. Royal Botanic Gardens, Kew. Pp. 285-333.

Whalen, M.D. 1984. Conspectus of species groups in Solanum subgenus Leptostemonum. Gentes Herb. 12:
179-282.

A NEW SPECIES OF CAMPOMANESIA (MYRTACEAE) FROM BAHIA, BRAZIL,
BASED ON SPECIMENS COLLECTED BY J.S. BLANCHE! OVER 150 YEARS AGO
Leslie R. Landrum Marla Ibrahim U. de Oliveira

School of Life Sciences
Arizona State University
Tempe, Arizona 85287-4501, U.l

Programa de Pds-Graduo(ao em Botdnica
Departamento de CUncias Bioldgicas
Universidade Estadual de Feira de Santana

Feira de Santana, Bahia, BRAZIL
marlauehbe@yahoo.com.br

ABSTRACT

RESUMO

Jacques Samuel Blanchet, Swiss ambassador in Bahia, Brazil, from 1828 to 1856 and an early plant collector
in that country (Staflue & Cowan 1976), has provided numerous type specimens, as can be deduced by
searching the epithets containing “blanchet” in the International Plant Name Index (http://www.ipni.org/
ipni/ plantnamesearchpage.do). Some genera are named in his honor, for instance Blanchetia DC. (Asteraceae)
(Stafleu & Cowan 1976). Most of these taxa were recognized as new in the 19‘*‘ Century, but some are still
being discovered based on Blanchet collections. For example, Landrum and Punch (2008) recently described
Psidium bahianum with a Blanchet collection cited as a paratype. The present new species is known to us by
only four sheets, one each at F and NY, and two at G, all distributed as Blanchet collections and possibly all
originally held at the Moricand Herbarium. This herbarium of ca. 50,000 specimens was amassed in the early
19* century by Moise-Etienne Moricand, a botanical associate of A. P. de Candolle (Stafleu & Cowan 1981).
The two G specimens have labels indicating that they were donated in 1908 by the grandson of Moricand.

Specimens of our new species have been filed as Psidium indet. and under “P. aggregatum,” a nomen
nudum, and Myrtus blanchetiana O. Berg (=P. salutare var. salutare; Landrum 2003) for perhaps more than
100 years. To even the trained eye, they would appear to be Psidium, but when an ovary was dissected,
the 8-10-locular ovary, with 7-10 ovules per locule, clearly placed the specimens in Campomanesia (Fig.
2D). Flowering specimens of Psidium and Campomanesia are superficially very similar and many species of
Campomanesia were originally described as Psidium. Fruit and seed characters separate the genera as well
(Landrum and Sharp 1989), but are not available for this new species.

The packet of the NY specimen contains a hand written note in Latin comparing the plant with species
of Psidium L. (some of which are now placed in Campomanesia Ruiz et Pav.) and Myrtus L. (one of which is
now placed in Calycolpus O. Berg). No species or genera of O. Berg are mentioned, so it is likely that the note
was written before the work of Berg on Brazilian Myrtaceae (1855-1859). Apparently Berg never saw any of
the sheets, as he doubtless would have described them as new. On one of the sheets from G there is a label
(in the same hand writing as the note at NY) identifying the locality of collection as Nazareth [Nazare] das
Farinhas and the collector as “M. Blanchet." Presumably, the note was not written by Blanchet, but could
have been written by Moricand.

I.Bot Res. Inst. Texas 4(2): 603 -6

Journal of the Botanical Research Institute of Texas 4(2)

Blanchet and “Nazare das Farinhas The collections attributed to Blanchet in Bahia extend from
Jacobina and Rio de Contas in the interior of Bahia to the coast (Urban 1840-1857). However, many of his
collections were made by others, and some Blanchet localities have still not been conhrmed. More confi-
dence can be placed in localities near the coast (A.M. Giulietti, pers. comm.). As indicated on one label,
Campomanesia blanchetiana was presumably collected at “Nazare das Farinhas” (commonly appearing on
maps simply as Nazare). Nazare is a center for the production of “mandioca” QAanihot esculenta Crantz.,
Euphorbiaceae) from which the food product “farinha” is produced, which accounts the town’s name. Since
the time of Blanchet this area, not far from Salvador, has been drastically altered by urbanization and the
cultivation of plantations of oil palm or “dende” (Elaeuisguineensis Jaquim, Arecaceae). Since our new species
has not been recollected in over 150 years the chances that it is now extinct are regrettably high.

Campomanesia blanchetiana Landrum & Ibrahim, sp. nov. (Fi^. 1, 2). Type: BRAZIL. Bahia: (fl) J.S. Blanchet 1611

Tree or shrub, glabrous except for minute hairs on some young growth (especially terminal buds), and in-
ner surface of calyx-lobes; hairs light reddish brown to whitish, less than 0.1 mm long; young twigs drying
reddish brown, the bark of older twigs light brown to yellowish, remaining smooth or somewhat flaky and
cracked; leaves elliptic to lanceolate, 3-10 cm long, 1. 2-4.2 cm wide, 2.4-3. 5 times as long as wide; apex
acuminate; base cuneate to acuminate; petiole 2-4 mm long, 0.5-1 mm wide, channeled; midvein impressed
above, raised prominently below; venation brochidodromous with ca. 12 lateral veins, these nearly straight,
leaving the midvein at an angle of more than 45 degrees, connecting to an arching lateral vein that follows
the margin, mostly within 1-2 mm of the margin; secondary veins dendritic, alternating with the laterals
and seeming to arise mainly from the marginal; blades subcoriaceous, drying reddish brown to yellowish
brown, densely glandular below. Flower buds broadly pyriform, to ca. 8 mm long, glabrous to minutely
puberulent, the base attenuate; peduncles 10-20 mm long, 0.6-1 mm wide, glabrous to puberulent, often
superimposed in pairs; bracteoles linear, caduceus before anthesis, ca. 1 mm long, ca. 0.2 mm wide; calyx
nearly closed in young bud, the terminal pore gradually expanding as the bud matures, with lobes reduced to
undulations along the margin, at anthesis tearing more or less regularly into 5 lobes, these broadly truncate,
ca. 1.5 mm long by 3-4 mm wide, the tears not entering the staminal ring; petals suborbicular to obovate,
7-8 mm long, glandular without; hypanthium infundibular, puberulent to glabrous, densely glandular; disk
(including staminal ring) ca. 5 mm across in recently opened flowers, glabrous; stamens ca. 10 mm long,
ca. 325; anthers ca. 0.5 mm long; style ca. 4.5 mm long, with a swollen base; ovary 8-10-locular; ovules
7-11; fruit unknown.

Additional specimens examined. BRAZIL. Bahia; “Nazareth [Nazarel das Farinhas," Blanchet 322 (G); “circa Bahia,” Blanchet 604 (NY).
’We believe that it is possible that all these Blanchet collections may be one gathering as they are all quite
similar, but since they have different numbers we cite them as separate collections. Although Blanchet 322 and
604 are technically paratypes, we believe that they should be preserved with the same care as isotypes.

Campomanesia blanchetiana is most similar to C. eugenioides and we compare the two in the key below.
1 . Calyx open in young flower bud, with lobes triangular, ovate, oblong, or hemiorbicular, 1-3 mm long and
wide, without tears forming between the lobes as the bud opens; lateral veins ca. 4-7 pairs, curving toward

apex; leaves mainly less than 2.5 times as long as wide; stamens up to ca. 2(X)

1 . Calyx nearly closed in young flower bud, with a terminal pore that gradually expands as the bud
with the lobes reduced to undulations along the margin in the closed bud, at anthesis tearing mo
regularly into 5 lobes, these broadly truncate, ca. 1.5 mm long by 3-4 mm wide; lateral veins ca.
nearly straight; leaves mainly more than 2.5 times as long as wide; stamens ca. 325

Fk. 2. Campomanesia blanchetiana, images of buds and flowers. A, B, C, flowers at various stages of anthesis; 1-4 indicate stages of opening, from
nearly dosed to after anthesis. D, aoss-section of ovary showing 8 locules, a typical number for Campomanesia. A, Blanchet 1611 at F; B & D, Blanchet
322atQ;C,Blanchet604.atm.

1 . Calyx closed in the bud, with an apiculate apex, at anthesis tearing irregularly; peduncles solitary; leaves 2-7

cm long, commonly with pocket-like folds of tissue on the lower surface (possible domatia) C. espiritosantensis

1 . Calyx nearly closed in young bud, with a terminal pore that gradually expands as the bud matures, at anthe-
sis tearing more or less regularly into 5 lobes; peduncles often superimposed in pairs; leaves 3-10 cm long
without folds of tissue on lower surface. C. blanchetiana

BOOK REVIEW

D. Jesse Wagstaff. 2008. International Poisonous Plant List: An Evidence-Based Reference. (ISBN 978-
1-420-06252-6, hbk.). CRC Press, Taylor and Francis Group, 6000 Broken Sound Parkway NW, Suite
300, Boca Raton, Florida 33487-2742, U.S.A. (Orders: orders@taylorandfrancis.com, 1-800-272-7737,
1-800-374-3401 fax). $149.95, 464 pp., 8 W x 11 1/4".

FOLIA TAXONOMICA 18. THE STATUS OF PASSIFLORA CITRIFOLIA AND A NEW
SPECIES IN SUBGENUS ASTROPHEA (PASSIFLORACEAE), PASSIFLORA JUSSIEUI
Christian Feuillet

Department of Botany
MRC-166, Smithsonian Institution
P.O. Box 37012

Washington, D.C 20013-7012, U.SA
feuillec@5i.edu

ABSTRACT

INTRODUCTION

Nomenclatural history of Passiflora citrifolia

The nomenclature for Passiflora citrifolia Ouss.) Mast., a species of subgenus Astrophea (DC.) Mast., has
evolved over the past 200 years. When Antoine Laurent de Jussieu (1805b, p. 392) described it in Tacsonia
Juss. (1789), he said that if this species were accepted, one could name it T. citriJoliaK He did not accept
that name, but merely proposed it for a future hypothetical naming whenever adequate material could be
studied. Jussieu (1805b) said about the material (herb. Richard) he studied that it was impossible to establish
a good species on such a specimen^ He stated that it was different from P. glandulosa Cav. and described
r. citrifolia as follows. It has oval entire leaves smooth and coriaceous, 13.5 x 8 cm (“5 x 3 pouces”) long,
petioles 5.4 cm long with 2 glands at the apex, not at the base. The peduncle and therefore the bracts are
missing. A detached flower has a tube nearly 4 cm long, a perianth divided in 10 oval lobes, a corona outer
row made of filaments a few lineae long (1 linea = 2.255 mm) [misspelled as “ligues” for lignes], another
inner row very small that seems to be made of glands. In the herbarium of the Museum d’Histoire naturelle
in Paris (P), there is a specimen clearly belonging to this species for the leaves, but with only small flower
buds, partly eaten by insects, in a pocket.

Augustin Pyrame de Candolle (1828, p. 335) validated the species by accepting it as T. citn/olia Juss.,
and the authorship should be given as Juss. ex DC. De Candolle placed Tacsonia citrifolia in Tacsonia section
Distephana DC. close to T. glandulosa (Cav.) Juss. and added that the leaves were oval, coriaceous, entire,
pinnately veined, and the petioles 2-glandular at apex “v. s. in h. Juss.” (= I have seen a dried specimen in
the Jussieu herbarium) without detail about the specimen. There is no material of P. citrifolia in the Jussieu
herbarium at P. Most likely de Candolle referred to the same material seen by Jussieu in 1805, and the de-
scription by de Candolle is shorter but agrees with Jussieu’s text.

Max Joseph Roemer (1846, p. 199) did not cite any specimen. He raised section Distephana to the rank
of genus and kept in the species that were in de Candolle’s section. For D. citrifolia Quss. ex DC.) M. Roem.,
he merely cited Jussieu and de Candolle, but the different brackets of the key leading to it could be patched
together into a description: leaves unlobed, entire, coriaceous, glabrous, acute(?), oval; petioles 2-glandular

610

the petiolar glands are basal versus apical and the bracts had basal glands when they were missing in the
specimen known to Jussieu, both points on which Jussieu (1805b) insisted. It strongly suggests that the
description was based on material representing P. glandulosa Cav.— his D. glandulosa (Cav.) M. Roem.

Maxwell Tyndell Masters (1871: 629) transferred the name to Passijlora L., unaware of, or dismissing, P
citrifolia Salisb. (1796). Richard Anthony Salisbury’s species is illegitimate because the original publication
included P. data Curtis 1788, a valid name, as a synonym (McNeill et al. 2006; ICBN art. 53.1). Nevertheless,
P citrifolia Salisb. (1796) has priority over P. citrifolia Quss. ex DC.) Mast. (1871), which therefore is illegiti-
mate as a later homonym (McNeill et al. 2006; ICBN art. 45.3). Later mentions of P citrifolia do not refer to
Salisbury’s name. Masters placed P. citrifolia (illeg.) in subg. Astrophea in his unnamed section 1 described
as: inflorescences cymose-paniculate, trees or shrubs often without tendrils. He cited “Sagot, 1287 &c.” as
a specimen.

Ellsworth Paine Killip (1938: 541) gave a good description of P. citrifolia and cited three specimens
from G, K, and P, all mentioned for the first time and collected later than 1805. He added: “none of these
specimens have good flowers,” but added that his description of the corona was based on “Sagot in 1857 (K)”.
In Paris (P) there is two sheets clearly labeled Sagot Oct. 1857 and in a thin folder there are two other Sagot
specimens, both with the mention “Herbier du Dr Sagot” and saying that it had been given by his widow in
1888; only one labeled Sagot N° 1287. Those specimens belong to the same species as the herb. Richard col-
lection. In Killip, the description of the leaves agrees with previous ones, except the one by Roemer. Killip’s
description is in conflict with Jussieu on the description of the flower with a shorter tube and a corona with
4-5 rows versus 2. There is doubt that the specimens Killip did study were of the same biological taxon as
the specimens examined by Jussieu and de Candolle, but they represent the species currently known from
the Guianas as P. citrifolia.

Different concepts of Passijlora citrifolia

The material seen by Jussieu was never cited clearly and had a flower in a bad condition according to Jussieu
(1805b) himself. In 1828, and de Candolle (1828) accepted Jussieu’s provisional name as a good species and
probably saw the same specimen. The collection used for the description apparently not seen since de Candolle
and the species designated by this name are not clear described the species currently called P. citrifolia.

Roemer (1846) studied different material and has a concept in conflict with Jussieu’s description. He
probably saw a small variation in what is now called P. glandulosa in subg. Passiflora supersect. Distephana
(DC.) Feuillet & J.M. MacDougal.

Masters (1871) did not give a description for the species and transferred it to Passiflora. Nevertheless,
through its placement in subg. Astrophea and the specimen cited, it is clear that the concept he had of the
species is different from the description of Roemer, but in agreement with Jussieu and de Candolle. Killip
(1938) described the flower from what is probably the same material and confirmed the placement in subg.
Astrophea.

The identity of the taxon described by Jussieu and accepted by de Candolle is uncertain due to the
poor quality of the potential type collection. A troubling fact is that Jussieu, de Candolle, and Roemer fail
to mention a clear character of the species currently known as P citrifolia— the leaves have a dark marginal
band when dried, an uncommon character in Passiflora. The modern concept of P. citrifolia does not match
Roemer’s description. Furthermore, it is not certain that the modern concept matches Jussieu’s comments
either. For those reasons, it seems better to describe the well documented species as a new species rather
than merely give it a new name.

Because the name Passiflora citrifolia Quss. ex DC.) Mast, is a later homonym, a other name is needed. In
the absence of synonym and in order to have for type a good blooming specimen, it seems better to describe
the well documented species as a new species rather than merely give it a new name.

Scandent shrub when young, then liana, glabrous throughout except the ovary; trunk woody, up to 3cm

red, 1. 5-4.5 cm long, adaxially 2-glandular at apex, the glands sessile, swollen, becoming saucer-shaped
when dry, yellow, when in growth, the young petiole oblique upward and the blade oblique downward dis-
playing the yellow glands as a crude egg mimic-structure; blades coriaceous, oblong or ovate-oblong, 12-20
X 5-12 cm, shortly acute and truncate or rounded at base, abruptly short-acuminate at apex, margin entire,
when dry there is abaxially a conspicuous marginal band, slightly recurved, narrow, dark brown or blackish
when dried, venation pinnate, main lateral veins 4-7(-9) each side of the midrib, strongly arcuate toward
the margin, prominently reticulate, when fresh pale green becoming dark with age, paler green abaxially.
Flowers solitary and axillary on young stems or in cauliflorous racemes with undeveloped leaves, racemes
often short, 1-5 cm long, sometimes as long as 50 cm, peduncles 4-6 mm long, bracts scale-like, glandless,
early deciduous. Flower tube cylindric, 1.5-2 cm long, white to green, often heavily marked with brownish
red; sepals 5, Ungulate, about 3x1 cm, spreading and often recurved along the tube, outside colored like
the tube, inside white; petals similar to the sepals in shape and habit, white; corona filaments in 4-5 series,
the outermost subdolabriform, spreading, 18 mm long, yellow-green, heavily marked with dark red except at
base and bright yellow apex, those of the next 2-3 series successively shorter, 1/3 to 1/6 as long, thick, oblique
to erect, colored like the first row, the innermost 1-2 mm long, filiform, some slightly capitate, reflexed into
the tube, white; operculum borne halfway up in the floral tube, slightly exerted, tubular, membranous at
base, laciniate at apex, white, light purple at apex; androgynophore 3-3.6 cm long, white to green; stamens
5, filament flattened, fused at base 1-2 mm, free part 7 mm long, white, anther rectangular, 6-7 x 2 mm,
pale yellow, pollen bright yellow; ovary narrow barrel-shaped, somewhat triangular in transverse section,
4-5 X 2-2.5 mm, strongly fluted when dry, densely short white- to green- or rufous-villous, 3-carpelled;
styles 7-9 mm long, diameter increasing from base to apex, pale green, stigmas capitate, 3 mm diam., yellow.
Fruit hanging, fusiform, hexagonal, seen only immature and green, glabrous; seeds not seen.

My above description agrees in most points with that given by Killip (1938, p. 541). Single axillary
flowers and cauliflory have been observed on the same plants. Those racemes are actually short stems with
bract-like leaves and either they continue normal vegetative growth, delayed or not, above the flowering
segment, or they end their growth and become caducous after fruiting. The same type of stems with a basal
inflorescence-like segment prolonged optionally by a normal leafy stem with axillary flowers is found toward
the base of the main stem in unrelated Passijlora species like, to cite a few, P. glandulosa Cav. (subg. Passiflora),
P. coriacea Juss. and P. suberosa L. (subg. Decaloba (DC.) Rchb.), or P. balbis Feuillet (subg. Astrophea).

Distribution and ecology. — Passiflora jussieui is known from French Guiana and Surinam in lowland
rainforest. It has been collected in bloom in Jan, Mar-July, and Sep-Oct. June is the rainiest month and
October the driest in French Guiana. It is likely that the species is not seasonal.

Etymology. — Passijlorajussieui has been named to honor Antoine Laurent de Jussieu (Juss.) who wrote
some of the early important papers for the understanding of the Passifloraceae (1789, 1805a, 1805b).

In cultivation. — Passijlorajussieui is one of the easiest species to propagate in a notoriously difficult sub-
genus. It was cultivated in the Botanical Garden of ORSTOM in Cayenne. That single plant is documented
by Cremers 6324 & 7156, Feuillet 1366, de Granville 5525 (all cited below) and was photographed several

Masters, M.T. 1871 . Contributions to the natural history of the Passifloraceae. Trans. Linn. Soc. 27:593-645.
McNeill J., F.R. Barrie, H.M. Burdet, V. Demoulin, D.L. Hawksworth, K. Marhold, D.H. Nicolson, J. Prado, P.C. Silva, J.E. Skog,
J.H.Wiersema, AND NJ.Turland 2006. International code of botanical nomenclature (Vienna Code). RegnumVeg.
146. A.R.G. GantnerVerlag KG, Ruggell.

Roemer, M.J. 1846. Passifloraceae. Prospect Fam. Nat. Syn. Monogr. 2:125-207.

Salisbury, R.C. 1796. Passiflora (154-156). In: Prodromus stirpium in horto ad chapel Allerton vigentium. s.n..

FOLIA TAXONOMICA 19. TYPIFICATIONS IN DILKEA (PASSIFLORACEAE)

Christian Feuillet

Department of Botany, MRC- 166
Smithsonian Institution
P.O.BOX37012

Washington, DC 20013-7012, U.SA

eDilkeahelleborifoliais

1 Dilkea Mast.

RESUME

sontd^signesetk

Dilkea johannesii Barb. Rodr., Vellosia ed.l, 1:24, pi. 10. 1888; ed. 2, 1:22 & 80 (caption); 3: t. 10. 1891.

Although not as well-known as the destruction of the herbarium in Berlin in 1943, the loss of the herbarium
of the Botanical Museum of Amazonia in Manaus had a serious impact on the nomenclature of Amazonian
plants. According to Mori and Castano Ferreira (1987), Joao Barbosa Rodrigues was the first and last director
of the Botanical Museum of Amazonia in Manaus. When he was nominated director of the Museum in Rio
de Janeiro in 1890, soon the Botanical Museum of Amazonia was closed and ultimately all the herbarium
specimens were lost. Among those, all Barbosa Rodrigues types collected before 1890 but one found later
in a book, were lost.

The first edition of Vellosia (Barbosa Rodrigues 1888) had so many errors that Barbosa Rodrigues
stopped the publication (Barbosa Rodrigues 1891a) and destroyed the copies at hand. The second edition
corrected most of the errors. The botanical illustrations (Barbosa Rodrigues 1891b) are reprints of Barbosa
Rodrigues original artwork, and in the case of D. johannesii, “tabula nostra X” cited above is the only original
element still extant. It is diagnostic with drawings and diagrams of the flower, fruit, and ovule. Therefore
I designate here tabula/estampa X (Fig. 1) as the lectotype of Dilkea Johannesii Barb. Rodr. 1888, dedicated
to his son joao Barbosa Rodrigues Jr.

An overlooked name in Dilkea

Barbosa Rodrigues (1891b) said that the genus Dilkea was described by Masters in “Flora Baziliensis” (sic) in
1872 with two species, D. retusa and D. acuminata, and two more added late, D. helleborijolia (as “hellibori-
folia”) and D. wallisii, and that he (Barb. Rodr.) is adding one more (D. johannesii, see above). In fact Masters
described Dilkea and the first two species in 1871. In “Flora Brasiliensis,” Masters (1872) gave a treatment
of the Passifloraceae where Dilkea is on columns 533-535 (columns, not pages are numbered), and Passi-
Jlora on 542-622. Then, also in 622, he described two species under “Curae posteriores,” Dilkea wallisii and
Passijlora helleborijolia (as “helliborifolia”) after illustrations from Wallis. Dilkea helleborijolia is probably an

on the genus. In any case, Passijlora helleborijolia Wallis ex Mast, and D. helleborijolia are synonyms of P. pedata
L. 1753 as noted by Killip (1938) for the former. As far as I know, nobody cited D. helleborijolia since 1891.

616

Journal of the Botanical Research Institute of Texas 4(2)

BOOK NOTICE

TAXONOMIC NOTES ON THE GENERA STENOTIS AND CARTERELLA
(RUBIACEAE) AND TRANSFER OF HEDYOTIS GREENEI TO STENOTIS

Edward E. Terrell

Harold Robinson

Department of Botany, MRC U
Smithsonian Institution
Washington, D.C 20013-7012, U.
Terr60@msn.com

Department of Botany, MRC 166
Smithsonian Institution
Washington, D.C. 20013-7012, U.S.A
Robinsoh@si.edu

ABSTRACT

s of some structures, geography, cytological c

RESUMEN

The genus Stenotis Terrell (2001) includes seven species restricted to Baja California and Arizona, two annu-
als and five perennial herbs or small shrubs. The present study is a reconsideration of taxonomic relation-
ships in the genus. Five of the species have chromosome numbers of n = 13 (Lewis 1962a), a number so far
known only in Stenotis and Carterella Terrell in the Hedyotis L./Houstonia L. relationship. A study by Church
(2003), using nuclear and chloroplast DNA sequences, included two species of Stenotis, S. arenaria and S.
asperuloides. and the single species, Carterella, and concluded that they were closely related to each other
but in a phylogenetic position outside of the Houstonia L./ Stenaria Terrell clade. The Stenotis clade remains
most distinct in its n = 13 chromosome number, and its near restriction to Baja California.

In our previous studies of Hedyotis/Houstonia relationship, we have emphasized the taxonomic impor-
tance of seed morphology. In the case of Stenotis the seeds of all species have punctiform hila on or near the
middle of the seeds. The annual species differ somewhat from the perennials in having the ventral, hilar
face somewhat enlarged and rounded (Table 1). Seeds of four species were shown by Terrell (2001, Fig. 1).

COMPARISON OF SPECIES

The present study is an addendum to the monograph of Stenotis by Terrell (2001), and a key to the species is
contained in that study. Six of the seven species of Stenotis are considered here. The present study does not
deal with S. peninsularis (Brandegee) Terrell, a very rare, twice-collected species. Of the remaining species,
we propose, based on our investigation, that one species in the previous treatment (Terrell 2001) is of hybrid
origin and Hedyotis greenei (A. Gray) W.H. Lewis be transferred from Hedyotis into Stenotis as discussed below,
a probable older name for the previously recognized Stenotis arenaria (Rose) Terrell.

Within Stenotis, the type species S. mucronata, is distinguished from all other species by having fas-
ciculate morphology, described as follows: Stem densely branched and with many short internodes, leaves

revolute, tightly clustered at the nodes in groups of 3-8 or more. This combination of characters gives the
plants a densely unique appearance. At least several species of Arcytophyllum Willd. ex Schult. in Roem. &
Schult., a related South and Central American genus, have similar fasciculate morphology; the relationships
of these species to Stenotis has not been tested, and is the subject of a separate study.

™ann.a, «

621

range south of La Paz in lower Baja California, whereas S. brevipes is rather widely distributed northward.
Terrell (2001) examined 13 herbarium specimens of S. australis, and 41 collections of S. brevipes.

The two annual species, S. arenaria (Rose) Terrell and S. asperuloides (Benth.) Terrell, occur in the Cape
region of Baja California south of La Paz. Both have short, slender herbaceous stems. When the two species
are compared with each other (Table 1) they differ as follows; Leaves slightly larger in S. arenaria and nar-
rowly oblanceolate to linear compared to linear or filiform in S. asperuloides. Pedicels in S. arenaria are to 16
mm long, compared to 30 in S. asperuloides. Corollas in S. arenaria 2-6 mm long vs. 3-1 1 mm. Corolla tubes
and lobes about equal in S. arenaria, but slightly longer in S. asperuloides. Capsules equally long and wide
in S. arenaria, but slightly longer than wide in S. asperuloides. Capsule shape in S. arenaria often subglobose
but often turbinate in S. asperuloides. Seeds are similar in both species. We conclude that the two annual
species have several differences, but are similar in important characters such as seeds.

Comparison of the annual S. arenaria with the perennial species having herbaceous stems, S. brevipes
(Table 1), shows differences besides those of plant size and life cycle: S. arenaria has corollas only 2-6 mm
long and corolla tubes about equal in length to the lobes, but S. brevipes has corollas 8-18 mm long with
tubes 3-4 times longer than lobes. Seeds of S. arenaria are slightly compressed and thick with the ventral face
strongly rounded. Seeds of S. brevipes are moderately compressed with both similar faces concave or convex.
These differences in reproductive characters suggest that these two species are more distantly related.

The other annual species, S. asperuloides, may also be compared (Table 1) with S. brevipes. Stenotis
asperuloides has pedicels to 30 mm long, corollas 3-11 mm long and corolla tubes only slightly longer than
the lobes. Stenotis brevipes has pedicels to 12 mm long, corollas 8-18 mm long, and tubes 3-4 times longer
than the lobes. The capsules of S. asperuloides are often turbinate and the length of capsules is longer than
its width. The capsules of S. brevipes are often subglobose and equally long as wide. Seeds of S. asperuloides
and S. brevipes differ as in S. arenaria and S. brevipes.

It was noted above that all three perennial species are closely related to each other in spite of the oc-
currence of fasciculate morphology in S. mucronata. The two annual species appear to be basically similar
to each other in important characters such as seeds, but in other characters they are distinct. It is concluded
that the annual and perennial species appear to be rather distinct groups, but are correctly placed in the
same genus, especially as no other genus except Carterella appears to be related to Stenotis.

ribed Houstonia gracilenta a
:r.” Wiggins (1980) stated tl
le species that Johnston coi

a San Diego Island (Moran 9592, ]

the type locality for S. gracilenta (Terrell 2001).

Stenotis gracilenta was tentatively treated as a species by Terrell (2001), who accompanied it with no-
menclatural data and a full description, but suggested that it is likely to be a hybrid between S. brevipes and
S. mucronata. Terrell (2001) provided a description to S. gracilenta and included it in the key to species. The

its hybrid status. It may be listed as Stenotis x gracilenta (I.M. Johnston) Terrell.

Transfer of Hedyotis greenii to genus Stenotis

Terrell (2001) stated that an additional species, Hedyotis greenei A. Gray, occurs only in Arizona and is closely
related to Stenotis arenaria, or the same species, but it was excluded from his treatment pending further study.
Since 2001 Terrell has not found time or opportunity for further study, however, data strongly suggest that
Hedyotis greenei is a Stenotis. It will take other studies to determine whether it is truly distinct from the very
morphologically similar Stenotis arenaria, but they are clearly both members of Stenotis. Whether it is distinct
from S. arenaria or is the same species or a variety remains to be determined. The geographical separation of
the two taxa— lower Baja California and Arizona amounts to ca. 960 km (600 miles). It may be noted that

623

anderae firmly within the Stenotis lineage. A question at the present time is whether Carterdla (1987) and
Stenotis (2001) are the same genus. Here the evidence is reexamined. Because Carterdla is in an unresolved
or basal position in the Stenotis clade (Church 2003), we suggest the lineage includes two genera, one being
the monophyletic Carterella.

east and southeast of La Paz in lower Baja California. It is quite different from the species of Stenotis discussed
here, and is a conspicuously attractive species with flower parts two to several times longer or larger than
those of Stenotis (Terrell 1987, Fig. 1). Its calyx lobes are 3-10 mm long versus 0.5-3 in Stenotis species;
corollas are 30-50 mm long versus 2-18; corolla tubes 25-41 mm long versus 1-13; corolla lobes 4.5-10
mm long versus 1-5; anthers 1.8-3 mm long versus 1-1.7; and stigma lobes 1-5 mm long versus 1-1.8. In
fruit the capsules length and width are 3-6 x 3-6 mm versus 1.3-5 x 1.3-4, significantly larger than most
if not all previously studied taxa in the Hedyotideae (s.s.). These measurements indicate that the length and
width of important flower and fruit characters are quantitatively different in Carterella.

Our previous work on the Hedyotideae (s.s.) found that seed morphology provided significant evidence
concerning the taxonomic status of genera and species. In studying Carterella alexanderae we found this
again to be true. Seeds of Carterdla alexanderae are strongly to moderately, laterally compressed with a hilum
at the peak (Terrell 1987, Figs. 2, 3) whereas seeds of Stenotis are somewhat dorsiventrally compressed, of
an ellipsoid type with a punctiform hilum on or near the center of the ventral face. Carterella alexanderae is
considered to be unique in having this strong lateral compression. Previous studies have not shown species
of Hedyotis/Houstonia relationship with such extreme morphology. Here it may also be noted that Bouvardia
seeds are dorsiventrally compressed with wide, papery, conspicuous wings and a chromosome number of 9.

basic stock as that of the Houstonia mucronata group” (now Stenotis), perhaps long isolated and the result

ng qual

)rphology. We 1

in flower and fruit
lat Carterella should

ACKNOWLEDGEMENTS

We thank Joseph Kirkbride for assistance with vagaries of computer procedure. We also thank the curators
of the cited herbaria for loans during past years and reviewers of the manuscript for providing helpful as-
sistance. Carlos Garcia-Robledo, Postdoctoral Fellow in the Department of Botany, is thanked for the Spanish
resumen. We thank David Lorence and an anonymous reviewer for helpful comments on an earlier draft.

Carter, A. 1955. A new species of Bouvardia (Rubiaceae) from Baja California, Mexico. Madrono 13:140-144.
Church, S.A. 2003. Molecular phylogenetics of Houstonia (Rubiaceae): descending aneuploidy and breeding
system evolution in the radiation of the lineage across North America. Molec. Phylogen. Evol. 27:223-238.
Johnston, I.M.1 924. Expedition of the California Academy of Sciences to the Gulf of California in 1 921 . Proc. Calif
Acad. Sci., ser.4, 12:1173-1177.

Lewis, W.H. 1 962a. Phylogenetic study of Hedyotis (Rubiaceae) in North America. Amer. J. Bot.49:855-865.

Lewis, W.H. 1962b. Chromosome numbers in North American Rubiaceae. Brittonia 14:285-290.

Lewis, W.H. 1968. Notes on Hedyotis (Rubiaceae) in North America. Ann. Missouri Bot. Card. 55:31-33.

Terrell E.E. 1987. Carterella (Rubiaceae), new genus from Baja California, Mexico. Brittonia 39:248-252.

Terrell E.E. 2001 . Stenotis (Rubiaceae), a new segregate genus from Baja California, Mexico. Sida 19:899-91 1 .
Terrell E.E. and H. Robinson. 2003. Survey of Asian and Pacific species of Hedyotis and Exallage (Rubiaceae) with
nomenclatural notes on Hedyof/s types. Taxon 52:775-782.

Wiggins, I.L. 1 980. Flora of Baja California. Stanford University Press, Stanford, CA.

BOOK NOTICE

Sabeeha Merchant, Winslow R. Briggs, and Donald Ort (eds). 2010. Annual Review of Plant Biology: Vol.
61 . (ISSN 1543-5008; ISBN 978-0-824-30661-8, hbk.). Annual Reviews, Inc., 4139 El Camino Way,
PO. Box 10139, Palo Alto, California 94303-0139, U.S.A. (Orders: www.AnnualReviews.org, science®
annualreviews.org, 1-800-523-8635, 1-650-493-4400). $89.00 indiv., 740 pp., 7%" x 93/8".

TRANSFER OF HEDYOTIS INTRICATA TO ARCYTOPHYLLUM (RUBIACEAE)

Edward E. Terrell

Research Associate
Department of Botany. MRC 166
Smithsonian Institution

Harold Robinson

Curator

Department of Botany, MRC 16

Washington. D.C 20013-7012. U

ABSTRACT

RESUMEN

Gray) Terrell & H. Rob. Se proporcioi

southern New Mexico, were studied. Morphological evidence indicates a close relationship to the genera
Stenotis and Arcytophyllum. After comparison of representative specimens of these genera, we conclude that
Arcytophyllum thymifolium (Ruiz & Pavon) Standi, is the most closely related species to Hedyotis intricata.
Transfer of Hedyotis intricata to Arcytophyllum, nomenclature, distribution, and representative collections
are provided below.

Distribution.— Rocky or gravelly slopes, arroyo banks, limestone ledges and crevices of cliffs, desert, semi-
desert, pine-oak, pine-juniper; 1000-2400 m. Mexico: Coahuila, western Nuevo Leon, east and central
Chihuahua, extreme north central Durango and Zacatecas; U.S.: southwestern Texas in southern Brewster
(including Chisos Mts., Big Bend National Park) and Presidio counties, and south central New Mexico (Dona

ACKNOWLEDGMENTS

We thank the herbaria cited in the text for loans during past years. We particularly thank Michael Powell,
director, SRSC herbarium, for loan of specimens from Brewster and Presidio counties, Texas.

REFERENCES

Mena V., P. 1990. A revision of the genus Arcytophyllum (Rubiaceae: Hedyotideae). Mem. New York Bot. Gard.
60:1-26.

Terrell, E.E. 2001 . Stenotis (Rubiaceae), a new segregate genus from Baja California, Mexico. Sida 1 9:899-91 1 .

NOMENCLATURAL TRANSFERS IN THE
GENUS MYRSINE (MYRSINACEAE) FOR NEW CALEDONIA
Jon M. Ricketson John J. Pipoly III

628

INTRODUCTION

The relationship among Myrsine L. and its satellites (inter alia Rapanea Aubl. and Suttonia A. Rich.) has been a
historical focal point for taxonomic contention that has resulted in the treatment of many species as Rapanea
by regional floristicians, while monographers have determined that the variation in androecial structure
among staminate flowers, and in gynoecial structure among the pistillate, supports the more broadly defined,
monophyletic concept of Myrsine. Regional monographers recognizing the more inclusive concept include
Hosaka (1940), Backer & Bakhuizen van den Brink (1965), Smith (1973), Fosberg & Sachet (1975, 1980),
Sachet (1975), Li (1978), Wagner et al. (1990), Pipoly & Chen (1995), Pipoly (1996, 2007), Jackes (2005)
and Takeuchi and Pipoly (2009) Among all of these papers, the most detailed treatment discussing all sides
of the issue is provided by Pipoly (1996) in treating Myrsine of the Philippines. Given all the reasons cited
in the aforementioned papers, we find that the recent new taxa and combinations made in Rapanea from
New Caledonia by M. Schmid (2009), need to be transferred to Myrsine, to provide a consistent treatment
of the genus pantropically, and necessitating the following new combinations:

Myrsine albiflorens (M. Schmid) Ricketson & Pipoly, comb. nov. Rapanea albijlorens M. Schmid, Adansonia, s6r. 3.
31:362. 2009, Type; NEW CALEDONIA: Kon6, Creek Pandanus, 21“02'04"S. 164‘>46'31"E, 27 Jul 2006 (fl). J. Munzinger & 1. Spir

1. Schmid) Ricketson & Pipoly, coml
E. NEW CALEDONIA; Haute vaUte de I’Amoa,
lez) Ricketson & Pipoly, comb.
e: new CALEDONIA; Piony, Sep 1868 (fr), B.
a (Mez) Ricketson & Pipoly st

Myrsine asymmetrica (Mez) Ricketson & Pipoly subsp. paniensis (M. Schmid) Ricketson & Pipoly, comb,
et stat. nov. Rapanea asymmetrica Mez var. paniensis M. Schmid, Adansonia, sir. 3. 31:350. 2009. Type: NEW CALEDONIA: Panii,
1200 m, 08 Apr 2006 (fr), J. Munzinger, Y Pilhn, H. Biajffart, M. Wanguene &■ I Spir 3468 (holotype: P; isotypes: MO, NOU).
Myrsine asymmetrica (Mez) Ricketson & Pipoly subsp. parvifolia (M. Schmid) Ricketson & Pipoly, comb, et
Creek Pernod, ca. 180 m, 10 Oct 1987 (fr), H. MacKee 43737 (holotype: P).

RicketsonandPipolyJ

hmid) Ricketson & Pipoly, comb. nov. s

)ore) Ricketson & Pipoly, comb.

1921. Type: NEW CALEDONIA: Montagnes au noid de la Ngoye,

Myrsine humboldtensis (M. Schmid) Ricketson & Pipoly, comb. nov. Rapanea hi

s6r. 3. 31:372, fig. 9A-D. 2009. Type: NEW CALEDONIA: Mont Humboldt, ca. 1500-1600 m. 23 Sep 1<

Myrsine kuebiniensis (M. Schmid) Ricketson & Pipoly, comb
31:366. 2009. Type: NEW CALEDONIA: Basse Ku^bini, ca. 5 m, 26 Nov
Myrsine lanceolata Pancher & Sebert, Not. Bois. Nouvel. Caledon. 192. 1874.
Mez, Pflanzenr. IV 236(Heft 9): 368. 1902. Type: NEW CALEDONIA: f '

descripti

Myrsine lanceolata Pancher & Sebert subsp.

CALEDONIA: Haute rivi&re des Pirogues, OuSnarou, ca. 150 m, 29 Aug 1987 (pist. f
Myrsine lecardii (Mez) Ricketson 6ar Pipoly, comb. nov. Rapanea kcardii Mez, pflanzenr. iv 236{Heft 9):369. i

Myrsine macrophylla (Mez) Ricketson & Pipoly, comb. nov. Rapanea macmphylla Mez, Pflanzenr. IV 236(Heft 9):369.

:c 1868 (fr), B. Balansa 992 (lectotype: P [designated

s (M. Schmid) Ricketson & Pipoly, comb.

ea macrophylla Mez var. menaziensis M. Schmid, Adansonia. ser. 3, 31:388. 2009. Type: NEW CALEDONIA:

(M. Schmid) Ricketson & Pipoly, comb. nov. Rapanea mcphersonii M. Schmid, Adansonia, ser.
2009. Type: NEW CALEDONIA: Sud: Mont Humboldt, 21°53'S, 166°25'E, ca. 1600 m, 19 Sep 1980 (fl), G.

Myrsine modesta (Mez) Ricketson & Pipoly, comb. nov. Rapanea modesta Mez, Pflanzenr. IV 236(Heft 9):3i
CALEDONIA: Au nord-est de la Conception, ca. 700 m, 07 Jan 1869 (fr), B. Balansa 991 (lecti
Myrsine modesta (Mez) Ricketson & Pipoly subsp. coriaria (M. Schmid) Ricketson & Pipoly, comb, et stat.
m, 13 Aug 1974 (fl, fr). H. MacKee 29119 (holotype: P).

Myrsine modesta (Mez) Ricketson & Pipoly subsp. tiebaghiensis (M. Schmid) Ricketson <Sr Pipoly, comb.

tiebaghiensis M. Schmid, Adansonia, ser. 3. 31:361. 2009. Type: NEW CALEDONIA; Partie
», 17 Oct 1969 (fr). H. MacKee 21016 (holotype: P).

kouC ca. 1250 m, 26 Nov 2002 (fr). J. Munzinj
ane nigricans (M. Schmid) Ricketson & Pipoly, comb. nov. Rapanea nigricans M. Schmid. A
fig. 4D-J. 2009. Type: NEW CALEDONIA: Mandj^lia, partie septentrionale de la chalne du Pani6 - Ignan
fl), H. MacKee 40580 (holotype; P; isotype; MO).

. Schmid) Ricketson & Pipoly subsp.

s (M. Schmid) Ricketson & Pipoly,

) Ricketson & Pipoly, comb. nov. Rapanea nmocaledonica Mez, Pflanzenr. IV 236(Heft
ONIA: bosque des environs de Eammia, Environs de Noumea, Feb 1871 (fl. fr), B. Balansa 3362

Myrsine novocaledonica (Mez) Ricketson & Pipoly subsp. balabioensis (M. Schmid) Ricketson & Pipoly,

Myrsine novocaledonica (Mez) Ricketson & Pipoly subsp. boulindaensis (M. Schmid) Ricketson & Pipoly,

LEDONIA; Noid: Poya Commune, Poya, mine Saint-Louis, ca. 20 m, 07 Sep 1969, H. MacKee 20693 (holotype: P).

s (M. Schmid) Ricketson & Pipoly,
s^r. 3. 31:354. 2009. Type: NEW CALE-
200 m, 18 Aug 1984 (fr). H. MacKee 42184 (holotype: P).

) Ricketson & Pipoly subsp. piroguensis (M. Schmid) Ricketson & Pipoly,

31:354. 2009. Type: NEW CALE-

s Pirogues, 15 Apr 1967 (fl, fr). H. MacKt

a (M. Schmid) Ricketson & Pipoly, c

:009. -frPE: NEW CALEDONIA: Bate de Prony, Sep 1868 (pist. D). B. B

631

Myrsine oblanceolata (M. Schmid) Ricketson & Pipoly subsp. doensis (M. Schmid) Ricketson & Pipoly,
DONIA: Mont Do. ca. 950 m. 28 Jul 1999 (pist. fl), T.Jaffri 3383 (hou)1ype: P; isotwes: NOU, MO).

Myrsine obovalifolia (M. Schmid) Ricketson & Pipoly, comb. nov. Rapmea obmahJoUa M. Schmid, Adansonia, s^r. 3.

I. Schmid) Ricketson & Pipoly, c

Letocart, R Amice & Catebk 3534 (holotwe; P; isotypes; MO, NOU).

Myrsine ouazangouensis (M. Schmid) Ricketson & Pipoly, comb

s6r. 3. 31:364, fig. 4K-M. 2009. Type: NEW CALEDONIA: Massif ultramafique du Ouazangou-T

Myrsine parvicarpa (M. Schmid) Ricketson & Pipoly subsp. amossensis (M. Schmid) Ricketson & Pipoly,
CALEDONIA: Col d'Amoss, ca. 450 m, 13 Nov 2002 (fr), J. Muminger et al. 1476 (holotype: P; isotype: NOU).

Myrsine parvicarpa (M. Schmid) Ricketson & Pipoly subsp. pachyphylla (M. Schmid) Ricketson & Pipoly,
om Kouakoue, 21“59'02"S, 166'>32'10"E, ca. 1100 m, 06 Nov 2004 (fr),J. LahatJ. Muminger &

Myrsine poumensis (M. Schmid) Ricketson & Pipoly, comb. nov. Rapaneapoumensis M. Schmid, Adansonia, ser. 3, 31:3
fig. 14D-E. 2009. Type: NEW CALEDONIA: Massif de Poum, ca. 350 m, 10 Sep 1'

Myrsine pronyensis (Guillaumin) Ricketson & Pipoly, com!

Myrsine taomensis (M. Schmid) Ricketson & Pipoly, coml
fig. 12E-0. 2009. Type: NEW CALEDONIA; Mont Taom, ca. 1000
Myrsine tchingouensis (M. Schmid) Ricketson & Pipoly,

I. 20‘’53'56"S, 165°00’27'E, 1260 m

Myrsine verrucosa (M. Schmid) Ricketson & Pipoly, comb. nov. Rapanea verrucosa M
figs. IE, 6A-D. 2009. Type: NEW CALEDONIA: Koniambo, 400-700 m. 21 May 1967 (fr), H. ,

Myrsine verrucosa (M. Schmid) Ricketson & Pipoly subsp. microphylla (M. Schmid) Ricketson & Pipoly,

632

HEDYCHIUM FORRESTII (ZINGIBERACEAE)

WITH A NEW SYNONYMY AND A VARIETY FROM INDIA
E. Sanoj,M.Sabu’andT.Rajesh Kumar

636

Journal of the Botanical Research Institute of Texas 4(2)

Distribution and ecology. — Hedychiumjorrestii var. palaniense is known only from four localities of Palani
Hills, Tamil Nadu at an altitude of 1300 -1600 m. It grows as patches of 15-20 mature individuals in grassy
slopes and open rock cliffs. It is assessed here as Critically Endangered (CR) (CRBlab(i,ii)+ 2ab(i,ii); D)
according to lUCN guidelines (lUCN 2001, lUCN Standards and Petitions Subcommittee 2010).

Flowering and fruiting.— July through October.

Etymology.— The varietal epithet has been derived from the type locality, i.e., ‘Palani Hills’, from where
the variety has been described.

to Sirur (near Ebanad), 6 Sep 1970, 1450 m,

2. Hedychium forrestii Diels, Notes Roy. Bot. Card. Edinburgh 5:304. 1912. (Fig. 2A). Type: CHINA. Yunnan:
Dali valley, 25°40'N, G. Forrest 4812 (E!).

Perennial rhizomatous herbs. Rhizome 3.3-3. 6 cm wide, creamy white internally, aromatic, covered by
brown scales. Leafy shoot 100-200 cm high, erect, robust. Leaves 12-18 in number, 12-16.2 cm apart,
sessile, spreading; sheath ca. 3.5 cm wide, green, margins pink, membranous, translucent, pubescent; ligule
3-5.2 X 2-2.3 cm, oblong, single lobed, slightly notched at tip, pale green, sericeous externally, membranous,
translucent, papery, closely appressed to the stem; lamina 34-55 x 6-13.3 cm, elliptic-lanceolate, dark green
and glabrous above (pink tinged in lower leaves), pale green and appressed silky hairy below; margin highly
undulate, membranous, translucent, pink tinged, non-ciliate, folded back; tip long caudate, twisted, bend
down; base oblique; midrib pink tinged below towards base. Inflorescence 15-27.6 cm long, cylindrical,
erect, moderately lax; peduncle green, pubescent. Bracts 4.8-5 x 2.5-2.T cm, obovate, green, pubescent,
dense towards tip and margins, leathery, convolute; margin non-ciliate, translucent, membranous, white;
tip rounded; cincinnus 2-5 -flowered. Bracteoles 3-3.7 x 2. 1-2. 3 cm, ovate, green, pubescent, outer one
tubular, inner ones non-tubular; margin membranous, translucent, non-ciliate. Flower 10-12.2 cm long,
white, spreading, 10-13 flowers opens at a time, ascending, slightly fragrant. Calyx 4.2-4.3 cm long, shorter
than bracts, 3.5-4 mm wide at mouth, white with pale yellow tinge, pubescent externally, membranous,
translucent, upper half inflated, lower half closely appressed to corolla tube, unilaterally split upto 1.2-1. 3
cm, 3-toothed at tip. Corolla tube 5-7.1 cm long, ca. 3.5 mm wide at mouth, white, glabrous externally,
downwardly directed hairy internally, glabrous towards base, erect or slightly bend towards tip. Corolla
lohes oblanceolate, white, pale green tinged towards tip, membranous, early decaying at base, drooping
from flower, glabrous, margins rolled inside, slightly coiled like an expanded spring; dorsal lobe 4.7-4.8
X ca. 0.6 cm, 5-6 mm long beaked at tip; lateral lobes 4.2-4.3 x ca. 0.5 cm, non-beaked at tip. Lateral
staminodes 3.4-3.7 x 0.8-1.4 cm, elliptic-oblanceolate, white, spreading on flower, reflexed back, tip
obtuse, margin undulate. Lahellum 3. 5-3.8 x 3. 1-3.5 cm, widely ovate-orbicular, white, pale yellow tinge
at centre and claw, spreading on flower, reflexed back on upper half, abruptly clawed at base; claw 6-8
mm wide; tip 1.2-1.7 cm deeply emarginated; margin undulate; lobes tip obtuse. Stamen 5-5.5 cm long,

0.25 cm, oblong, creamy white, split opens from top to bottom, attached with the filament at ca. 2.5 mm
above from base, thecae parallel with the filament axis; connective white, glabrous, non-crested, slightly
notched, tip truncate. Ovary 4.5-5 x 3-3.5 mm, barrel-shaped, pale green, densely pubescent, trilocular.
ovules many, placentation axile. Style filiform, white, glabrous, pubescent towards tip, broadens and green
tinged towards stigma. Stigma green, cup-shaped, slanting with a depression at center, densely pubescent,
hairs hook-like, bulbous based, 1-1.5 mm exserted from the anther. Epigynous glands two, 3-3.5 mm
long, oblong, bright yellow. Fruits 4-6 x ca. 1.5 cm, oblong-cylindrical.

637

Type specimen (f.SflTOyS6;57).

638

Journal of the Botanical Research Institute of Texas 4(2)

Notes. — Hedychiumforrestii was originally described by Diels in 1912, based on a collection by Forrest
(holotype; G. Forrest 4812, E!) from Tali valley in Western Yunnan. He distinguished this species from H.

staminodes.” However, he admitted that, he was not able to study some of the floral details of this species.
Diels’ diagnosis of this species was rather short and was based on a specimen with a single upper leaf and
an inflorescence. Later, while describing the new variety from Vietnam (var. latebracteatum), Larsen (1965)
elaborated on the description of H.forrestii based on his studies at Kew, Paris, and Edinburgh.

In the meanwhile, Rao and Verma (1969) apparently being unaware of the description of H.forrestii,
described yet another species, Hedychium dekianum from Jowai, Assam (presently in Meghalaya), based on
a collection by Deka (C.K. Deka 35605 A). The authors compared their new species with H. coronarium and
distinguished it by its “lax spikes; closely convolute bracts; stamen exceeding the lip and oblong-cylindric

During the course of revision of the genus in India, we were able to collect and study a large number
of specimens of Hedychium from the entire Northeast India. Our studies on H. dekianum based on the types
and live specimens have indicated that Indian specimens differ from H. forrestii in the white color of the
filament and creamy white anther, whereas it is salmon pink and crimson respectively in H.forrestii, which
is mentioned only in one of the collections of G. Forrest 8478 (K!). As the differences between these taxa are
not sufficient to recognize them even as varieties, H. dekianum is reduced to a synonym of H.forrestii.

Distribution and ecology. — Distributed in Yunnan and Guizhou provinces of China, India, northern
Thailand, Myanmar, and Vietnam. In India it is restricted to the districts. East Khasi Hills and Jaintia Hills
of Meghalaya. It is a robust herbaceous plant growing in populations of 3-5 plants in open and moist forest

Hedychium forrestii is facing a very high risk of extinction in the wild. Hence we categorize it as Endan-
gered (EN) (CRBlab(i,ii,iv)+ 2ab(i,ii,iv) as per lUCN guidelines (lUCN 2001, lUCN Standards and Petitions
Subcommittee 2010).

Council of Scientific and Industrial Research (CSIR), New Delhi for the award of Senior Research Fellowship
to the first author (9/43(0138)2K9-EMR I), and the research grant provided by the Department of Science and
Technology. Govt, of India. New Delhi (SP/SO/PS-115/2009) are gratefully acknowledged. We are thankful to
J.F. Veldkamp, Leiden, The Netherlands for the Latin diagnosis. We are also grateful to the Director, Botani-
cal Survey of India, and curators of BM, E and K for the kind permission for consultation of their Herbaria.
The digital image of H.forrestii type provided by E is greatly acknowledged. Various help rendered by A.K.
Pradeep, Curator, Calicut University Herbarium (CALI) for the preparation of manuscript is also gratefully
acknowledged. Mark Newman and an anonymous reviewer provided constructive reviews.

REFERENCES

Diels, L. 1912. Plantae Chinenses Forrestianae. Notes Royal Bot. Gard. Edinburgh 5:304.

Jain, S.K. and V. Prakash. 1 995. Zingiberaceae in India: phytogeography and endemism. Rheedea 5:1 54-169.
Koenig, J.F. 1783. Observationes botanicae (Retzius) 3:73-74.

BOOK REVIEW

Toutcha Lebgue Keleng and Gustavo Quintana Martinez. 2010. Cactaceas de Chihuahua: Tesoro estatal en
peligro de extincion. (ISBN 978-6-077-78834-8, pbk.). Published by the authors. (Orders: Contact
Robert Armijo at: rarmijp@yahoo.com; accepting checks only). $25.00, 248 pp., color photos, b&w
maps, 5Ya” x 8ys".

THE REINSTATEMENT OF PTILIMNIUM TEXENSE (APIACEAE)
AND A NEW KEY TO THE GENUS

Mary Ann E. Feist

//linois Natural History Survey
University of Illinois at Urbana-Champaign
1816S.OakSt.

ABSTRACT

RESUMEN

Ptilimnium texense J.M. Coult. & Rose is currently most commonly treated as a synonym of P. costatum (Elliott)
Raf., but some authors continue to recognize it as the hybrid P. xtexense, with the putative parents P. capil-
laceum (Michx.) Raf. and P. nuttallii (DC.) Britton. A recent study utilizing nrDNA ITS sequences indicated
that Ptilimnium texense was more closely related to P. costatum than to P. capiUaceum or P. nuttallii (Feist &
Downie 2008), but also brought into question the synonymy of P. texense and P. costatum. This was followed
by extensive herbarium and field studies to examine the taxonomic status of P. texense. Combined evidence
from morphology, ecology, and DNA sequences indicates that P. texense should be reinstated. Ptilimnium
texense occurs in acidic habitats in the West Gulf Coastal Plain Region of east central Texas, west central
Louisiana, and southern Arkansas (Fig. 1).

Taxonomic History. — In 1909 J.M. Coulter andJ.N. Rose described Ptilimnium texense based on a speci-
men from Hockley, Texas, collected by F.W. Thurow. They noted that it combined “the cleft involucral bracts,
characteristic fruit ribs, and shorter styles of P. capiUaceum with the stouter habit, smaller fruit, and larger
calyx teeth of P. nuttallii.” In 1945, Mathias and Constance speculated that plants described as P. texense
“seemed to be of hybrid origin,” and listed the putative parents as P. capiUaceum and P. nuttallii. Easterly
(1957) was unable to find distinctive characters for P. texense and so accepted this assessment, stating that
“this plant combines the fruit characteristics of P. nuttallii with the vegetative characteristics of P. capiUaceum.”
None of these authors, however, presented evidence beyond these statements to support the hybrid status of
P. texense and later Mathias and Constance relegated it to synonymy under P. costatum with no explanation
(Lundell 1961). Some authors, however, have continued to recognize P. xtexense (Correll & Johnston 1970;
Correll & Correll 1972; Diggs et al. 1999) and have given its distribution as eastern Texas in acid bogs and
marshlands. These same authors have also continued to recognize P. costatum as occurring in Texas.

Molecular Studies.— In a study utilizing nrDNA ITS sequence data (Feist & Downie 2008), specimens

642

lentified by Easterly as P. xte
laceum and P. nuttallii), but rather showed
strongly supported clade with all specimen

ith P. costatum. These specimens formed a
urn from Texas and Louisiana (hereafter the
R texense clade). The R texense clade was sister to another strongly supported clade made up of R costatum
from Georgia, Illinois, Missouri, and Tennessee (hereafter the R costatum clade). These results indicated that
R xtexense was allied with R costatum, but that populations of each from Louisiana or Texas (the R texense
clade) were molecularly distinct (Feist & Downie 2008). Average ITS sequence divergence between the R
texense and P. costatum clades was 2.6%, which is just slightly less than that between R costatum and R capil-
laceum (2.9%). Results from a recent analysis of cpDNA sequence data (trnQ-rpsl6 5’exon, rpsl6 intron, rpsl6
3’exon-trnK) were congruent with the ITS findings (Feist & Downie unpublished data). At first glance, these
results seemed to support Mathias and Constance’s conclusion (1961) that P. texense should be synonymized
under R costatum, but morphological differences between specimens making up the R texense clade and the
P. costatum clade were also observed. Taken together with the geographical and molecular differences, this
suggested that members of the R texense clade might represent a taxon distinct from P. costatum. Additional
herbarium and field studies were undertaken to investigate the taxonomic status of these populate
Morphology, habit, phenology, habitat requirements.

1 considered. The results of

METHODS

Field Visits and Morphological Studies. — Six populations of Ptilimnium, three in Louisiana and three in Texas,
all previously identified as P. costatum but determined to be P. texense during the course of this study, were
visited during Sep 1 5-2 1 , 2009 (Feist & Molano-Flores specimens listed in Appendix 1). Habitat and associate
species were recorded and voucher specimens were collected for additional morphological study. In addition,
a total of five plants were collected live and placed in a greenhouse at the Illinois Natural History Survey.

A large number of specimens from the genus Ptilimnium were examined during the course of this study.
Specimens were either collected by the author or borrowed from the following herbaria: ANHC, AUA, BAYLU,
BRIT, DOV, DUKE, EKY, F, FLAS, FSU, GA, ILL, ILLS, JEPS, LAF, LL, LSU, LSUS, MO, NCSC, NCU, NO, NY,
OKL, OKLA, OS, OSC, PH, RM, RSA-POM, SMU, TAMU, TENN, TEX, UARK, UC, UNA, UNC, US, USCH,
USF, USFS, and WVA. A total of 144 specimens were determined to be Ptilimnium texense. These included 4
from Arkansas, 54 from Louisiana, and 86 from Texas (Appendix 1). The majority of these specimens were
not identified as P. texense prior to this study, but were annotated to P. texense by the author.

Morphological data were collected from herbarium specimens. These are summarized in the taxonomic
description below, and the characters critical for species identification are highlighted in the key to Ptilimnium.
Phenology, habitat, and distribution were determined from the herbarium specimens and the accompanying

field and in the greenhouse.

HABITAT AND DISTRIBUTION

According to the USDA Plants Database, which treats Ptilimnium texense as a synonym of P. costatum, P. costatum
occurs in 25 parishes in Louisiana and 13 counties in Texas (USDA. NRCS 2010). All specimens labeled as
P. costatum examined by the author from these states are P. texense (10 parishes in Louisiana and 19 counties
in Texas) or were misidentified collections of P. capillaceum or P. nuttallti. No specimens of P. costatum from
Texas or Louisiana were found. Accessions of P. costatum cited from Dallas and Grayson counties (Mathias
& Constance 1961; Diggs et al. 2006) could not be located. Both species grow in Arkansas but are allopatric,
with P. texense occurring in two southern counties and P. costatum in three counties farther north (Fig.l).

Ptilimnium texense is endemic to the West Gulf Coastal Plain (WGCP). This physiographic region encom-
passes much of eastern Texas, western Louisiana, southeastern Oklahoma, and southern Arkansas (Fig. 1).
Approximately 3900 species occur in the WGCP, which has been divided into four ecoregions: Oak-Pine-
Hickory Forest, Longleaf-Pine Forest, Post Oak Savanna, and Prairie (MacRoberts & MacRoberts 2003).
Ptilimnium texense occurs primarily in the Oak-Pine-Hickory Forest and Longleaf-Pine Forest ecoregions.
The primary habitats of P. texense are bogs, acid seeps, and wet pine savannas. Common associates include
Pinus palustris, Liquidambar styraciflua. Acer rubrum, Nyssa syhatica, Magnolia virginiana, Sphagnum spp., Sar-
racenia alata, Oxypolis rigidior, Eryngium integrifolium, Eriocaulon spp., Osmunda regalis and O. cinnamomea.
Ptilimnium texense can be added to the list of 96 species endemic to the WGCP (MacRoberts et al. 2002).
About 9% of WGCP endemics are found in bogs/wet pine savannas, which occupy 10% of the total area of
the WGCP (MacRoberts et al. 2002).

DISCUSSION

Ptilimnium texense was proposed as a hybrid of P. capillaceum and P. nuttallti because earlier authors felt that
P. texense combined characteristics of these species and possessed no stable characters of its own (Mathias
& Constance 1945; Easterly 1957). Upon closer inspection of additional specimens, unique characters that
distinguish P. texense were found. Contrary to Easterly’s assessment, the fruits of P. texense and P. nuttallti
are easily differentiated. The fruits of P. texense are longer than those of P. nuttallti (2.2-3.5 mm versus 1-1.9
mm) and the dorsal ribs are narrow and blunt versus thick and rounded. Vegetatively, P. texense differs from
P. capillaceum in that its leaf segments are always 3-4-angled to subterete and the midveins are not visible.

644 Journal of the Botanical Research Institute of Texas 4(2)

whereas the leaf segments of P. capillaceum are often flattened and the midveins are apparent. Unique char-
acters of P. texense not found in P. capillaceum or P. nuttallii include root system and habit. Both P. capillaceum
and P. nuttallii have fibrous roots and are annuals, whereas P. texense has a distinctive rounded corm at the
base of the stem and is a perennial. Ptilimnium texense has a more limited geographic range than either P.
capillaceum or P. nuttallii, and more restrictive habitat requirements, as it requires acidic environments such
as bogs, acid seeps, and wet pine savannas. Where the ranges of the three species overlap, P. capillaceum and
P. nuttallii flower and fruit much earlier than P. texense. By the time P. texense begins to flower in August, the
fruit of P. capillaceum and P. nuttallii have already matured and fallen from the plant. The fruit of P. texense

Molecular data provided no evidence that P. texense might be a hybrid. The results of studies utilizing
nuclear and chloroplast DNA sequences were congruent (Feist & Downie 2008; Feist & Downie unpublished
data) and there were no site polymorphisms visible on the chromatograms of accessions identified as P. texense.
Incongruence and site polymorphisms may both be signs of hybridization (Baldwin et al. 1995; Alvarez &
Wendel 2003). Taken together with the lack of morphological evidence provided by earlier authors and the
distinctive characters of P. texense presented in this study, the hybrid status of P. texense is not supported.

Ptilimnium texense was synonymized under P. costatum (Mathias & Constance 1961) and it is most closely
related to this species (Feist & Downie 2008). These species share some morphological traits, such as root
morphology and perennial habit, but are also distinct in a number of ways (Figs. 2, 3). These differences
are highlighted in the last two couplets of the key. A major difference distinguishing the species is their leaf
morphology. As with P. capillaceum, leaf segments of P. costatum differ from those of P. texense by being flat
with the midvein apparent. Other differences in leaf morphology include geometry and overall shape. The
leaves of P. costatum are distinctive in that the primary leaf segments become progressively shorter from the
leaf base to the apex and the blade apex is acute. In P. texense, the primary segments remain about the same
length along the rachis and the blade apex is blunt or rounded. The leaves of P. texense are also suffer and
more three-dimensional than the leaves of P. costatum and the other Ptilimnium species. Style length also
distinguishes P. texense and P. costatum. Although there can be some overlap, the styles of P. costatum are typi-
cally longer [(0.8-)l-2 mm] than those of P. texense [(0.3-)0.5-0.8(-l) mm]. There is no overlap, however, in
the geographic range of these species and their habitats are different as well, with P. texense again requiring
a more acidic environment. As mentioned above, both P. costatum and P. texense are perennials with corms.
This structure is usually more elongate in P. costatum than in P. texense and may be slightly L-shaped. These
characters have not been noted in previous studies of Ptilimnium. All species of Ptilimnium had been thought
to be annuals with fibrous roots. P. costatum and P. texense do develop numerous adventitious roots which
may obscure the corm. but both species develop buds from the corm (cormlets) that grow into new stems
(Fig. 4).

As P. texense has been purported to be a hybrid or not differentiated from P. costatum, it has not been
included in keys to the genus. It clearly does not fit the description of any of the other species and this has
lead to confusion when attempting to identify it. Of the 138 specimens identified as P. texense in the course
of this study (not including the author’s own collections), 22 were not initially identified to species or not
identified as Ptilimnium, 34 were identified as P. capillaceum, 44 as P. costatum, 36 as P. nuttallii, and just

costatum by subsequent workers. This poor record of identification clearly demonstrates the need for a more
inclusive and refined key. Previous keys to the genus have emphasized style length, number of segments
of the involucral bracts, and whether the primary leaf segments are alternate or opposite on the rachis. Al-
though these characters can be useful, there is variability and overlap among them and relying exclusively
on these characters can lead to confusion. For this reason, additional characters have been utilized in the
key below.

edbyDi.Correllin1%9.

R.Phillippein1993.

647

648

KEY TO PJILIMNIUM

. Middle and upper petiole bases papillate on the abaxial surface; styles 0.1 -0.2 mm long (on fruit),

ascending to spreading; calyx teeth < 0.2 mm, deltoid P. c

. Middle and upper petiole bases not papillate on the abaxial surface; styles (0.3-)0.4-2 mm long (on fruit),
spreading to strongly recurved; calyx teeth > 0.2 mm, narrowly triangular.

2. Mid-stem leaves with 2-4(-5) nodes along the rachis, primary leaf segments usually alternate or opposite
at the nodes (not including at the apex of the petiole); individual leaf segments often much longer than
the rachis; involucral bract segments 1 (-3); roots fibrous, stem sometimes slightly thickened at the base
but never forming a corm; styles (0.3-)0.4-0.6 mm long; fruit 1-1.9 mm long, dorsal ribs thick, rounded;

flowering Apr-Jul, fruiting late May-early-Aug

2. Mid-stem leaves with (6-)7-16 nodes along the rachis, primary leaf segments whorled or verticillate at
the major nodes; individual leaf segments shorter than the rachis (rarely as long as in P. texense): involucral
bract segments (1 -)3(-7); stem thickened and rounded at the base forming a globose or slightly elongate
corm; styles (0.3-)05-2 mm long; fruit 2.2-4 mm long, dorsal ribs narrow, sharp-edged to blunt; flowering
Jul-Oct, fruiting mid-Jul-Nov.

3. Leaf segments flat (at least near the nodes but usually throughout), midvein visible; mid-stem leaves
with (8-) 10-1 6 nodes per rachis, longest primary segments with 9-22 secondary segments; leaf shape
in outline deltoid or trullate (trowel-shaped), leaf segments getting progressively and noticeably
shorter towards apex, apex acute; styles (0.8-)1 -2 mm long, slightly spreading to spreading; flowering
Jul-Oct, fruiting mid-Jul-Oct; bottomland forest, swamps, streambanks, and pond margins

3. Leaf segments 3-4-angled to subterete, midvein not visible; mid-stem leaves with (6-)7-1 0 nodes per
rachis, longest primary segments with 3-7(-8) secondary segments; leaf shape in outline oblong or
oval, leaf segments not getting progressively and noticeably shorter towards apex, apex blunt; styles
(0.3-)0.5-0.8(-1 ) mm long, spreading to strongly recurved; flowering Aug-Oct, fruiting Sep-Nov; seeps.

lants perennial, 5-12 dm, roots f
tiff, 3-dimensional, oblong to oval i
ichis, 3-5 primary leaf segments at the major nodes; leaf segments
5 subterete, midvein not visible, individual leaf segments shortei

stem. Leaves: blades 3-12 cm,
s with (6-)7-10 nodes along the
:d, filiform to linear, 3-4-angled
-as long as) the rachis, primary

leaf segments not getting progressively and noticeably shorter towards apex, longest primary segments with
3-7(-8) secondary segments; petioles 0.5-3 cm, hyaline borders narrow, often inconspicuous and only vis-
ible at the distal and proximal ends, abaxial surface not papillate at the base. Peduncle 2-12 cm. Umbels
3-8(-10), rays 8-20(-25), l-3.5(-4.5) cm, subequal to of varying lengths; involucral bracts linear, entire
or 3-parted, rarely further divided. Pedicels 2-12 mm. Flowers 12-24 per umbellet; calyx teeth conspicu-
ous, 0.2-0.4 mm, narrowly triangular; petals (0.6-)0.7-1.3 mm; styles (0.3-)0.5-0.8 (-1) mm, spreading
to strongly recurved. Schizocarps 2. 2-3. 5 x 1.5-2. 2 mm, ovate to orbicular, slightly compressed laterally,
often maroon- or purple-tinged, dorsal ribs narrow, blunt, corky-thickened extension of the lateral ribs
conspicuous; oil tubes dark brown. Flowering/fruiting Aug-Oct/Sep-Nov.

NO). Columbia Co,: Emerson, 8 Sep 1 948, D.M. Moore 480649 (5MU, UARK).

LOUISIANA. Allen Pa.: off LA 1 12 ca. 4 mi W of Elizabeth, infrequent in pine forest, 5 Sep 1981, C.M. Allen 11315 (LSU). Beau-
regard Pa.: beside LA 1 10 6.5 mi SE of Merryville near Bearhead Creek, wet area, 3 Aug 1974, R.D. Thomas 40979 (TENN).
Calcasieu Pa.: Lake Charles, prairies, 1 4 Sep 1 906, R.S. Cocks s.n. (NO); 4 mi S of Holmwood, grassy roadside, 23 Sep 1 967, J.W.

650

E. Larson Sandylands Preserve, Tract 4, ca, 33 tmi E of jet. of Hwy. 69 and Hwy. 418 on 41 8, S side of 41 8 and W side of Village
Creek, 1 2 Aug 1 994,i Singhurst 2270 (BAYLU); Roy E. Larson Sandylands Preserve, Tract 4, ca. 3.3 mi E of jet. of Hwy. 69 and Hwy.
41 8 on 41 8, S side of 41 8 and W side ofVillage Creek, 1 5 Oct 1 994, J. Singhurst 20 1 9 (BAYLU); Roy E, Larson Sandylands Preserve,
Tract 4, ca. 3.3 mi E of jet. of Hwy. 69 and Hwy. 41 8 on 41 8, S side of 41 8 and W side ofVillage Creek, 1 5 Oct 1 994, J. Singhurst
2050 (BAYLU); roadside ditch just outside of Roy E. Larson Sandylands Preserve, within the Hyatt Lake Estates, roadside ditch
near longleaf pine savannah, 25 Sep 2009, M.A. Feist &B. Molano-Flores 4464 (ILLS). Harris Co.: near Hockley, Sep 1890, F.W.
Thurows.n. (US). Henderson Co.: 8 mi from Athens, seepage slope along stream, 28 Aug 1946, V.L Cory 14188 (LL). Houston
Co.: Grapeland, open sandy bogs, 1 6 Sep 1 91 8, EJ. Palmer 14423 (MO). Jasper Co.: off TX 63 NW of Beans Place, in pineland,
10 Sep 1942, CL&A.A. Lundell 1 1903 (LL); in longleaf pine region in rolling country, 6 mi N of Kirbyville, evergreen shrub bog,
1 0 Nov 1 962, D.S. Correll 26755 (LL); 2 mi W of McGee Bend Dam, seepage slope among Magnolia virginiana and Sarracenia, 1 3
Nov 1 963, D.S. Correll 28638 (LL); Boykin Bog, bog, 23 Aug 1 964, R.P. Turner 144 (TEX); N side of Little Rocky Creek NE of Grubbs
House, Little Rocky Preserve, ca. 3.4 mi E of jet. US 96 and FM 1 007 near Browndell, locally frequent in wet sandy soil in forested
seep in loblolly pine-hardwoods forest below open hillside seepage bog, 24 Sep 1997, W.R. Carr J 7060 (TEX); E of CR 348A,
along small roadside stream in cutover longleaf pine savannah, 25 Sep 2009, M.A. Feist &B. Molano-Flores 4458 (ILLS); E of CR
348A, along small stream in cutover longleaf pine savannah, 25 Sep 2009, M.A. Feist 8iB. Molano-Flores 4463 (ILLS). Jefferson
Co.: Beaumont, 1 5 Sep 1 936, J.L Flooks s.n. (TEX). Leon Co.: ca. 8.5 mi NE of Buffalo on US 79, seepage Sarracenia bog, 1 6 Sep
1 994, W.C. Holmes 7466 (BAYLU); Cripple Fawn Ranch, NE of Flynn, Sarracenia bog with Pluchea, Myrica, Lobelia, and Solidago,
10 Nov 1995, M. DubruleReed 1738 (TAMU). Nacogdoches Co.: near Nacogdoches, 24 Sep 1938, £ Whitehouse 1 1376 (SMU).
Newton Co.: SF 1, 5 mi E of Kirbyville, moist sandy woods, 30 Sep 1945, V.L. Cory 4981 7 {SMU, US); 1.5 mi E of county line
along Farm Rd. 363 to Bon Wier, savannah between road and railroad, 30 Oct 1 968, D.S Correll & H.B Correll 36692 (LL); 5 mi
due W of Deweyville, edge of moist woods, 1 4 Sep 1 968, D.S. Correll 36530 (LL); Scrappin Valley Distinctive Site, Temple 1 niand
Corp Land, 5. 1 mi N of jet. of R 255 and Hwy. 87 at Mayflower Community on Hwy. 87, E side of Hwy. 87, Sphagnum-beakrush
community, hill-side seepage slope bogs, acid seep spring bogs, 1 5 Oa 2002,1 Singhurst 1 1262 (BAYLU). Robertson Co.: near
Newbaden, widespread on bog, 30 Oct 1 943, JJ. Brady, B.C. Tharp, &F.A. Barkley ?3750(DUKE, MO NO, OKL, OKLA, RM,TEX, UARK,
UNC, US); 1 5 mi SE of New Baden, moist area of bog, 1 1 Oct 1 948, EM. Trew, Jr. % (TEX); 4 mi E of New Baden, bog, 2 1 Aug
1948, G.L Webster & CM. Rowell 1 904 (TEX); 5 mi E of New Baden, peat bog, 24 Oct 1 948, G.L. Webster & CM. Rowell 1953 (TEX,
UARK); Southworth bog, ca. 1 5 mi SE of New Baden, moist portion of bog, 1 1 Oa 1 948, 11 Sperry 2 157 (TAMU); Southworth
Peat Bog, 1 2 mi E of New Baden, bog in Carrizo sands with cattails, pitcher plants, and panic grass, 20 Aug 1 954, EL Rabb 108
(TAMU); S.W. 1/4 Camp Creek Lake Quad., Mill Creek Bog, bog, 4 Oct 1981, T Starbuck 1097 (BRIT, TAMU). Smith Co.: Swan,
swamps, 1 7 Sep 1902, 1 Reverehon 3193 (MO, US); Swan, swamps, 1 7 Sep 1902, 1 Reverchon s.n. (MO, US); western Tyler, on
ROW of Cotton Belt Railway, frequent in muddy ground along a small drainageway, 5 Sep 1 949, V.L. Cory 56892 (US). Tyler
Co.: Kirby Forest, near Warren and Kountze, 25 Aug 1 945, S.R. Warner 475 (TEX); ca, 2 mi NE of Warren, in a draw near a creek,
1 Oa 1 945, V.L Cory 49869 (OKLA, SMU, US); 25 mi S of Warren, frequent in pitcher plant bog, 28 Sep 1 948, V.L Cory 54898 (LL,
SMU); ca. 1 0 mi E of Hillister on road to Spurger, evergreen shrub bog, pinelands, 1 5 Nov 1 963, D.S. Correll 28680 (LL); ca. 2 mi
S of Warren on Rte. 69, boggy area, 1 5 Oct 1 969, D.S. Correll 38163 (LAF, LL, MO, UNC); Turkey Creek Unit, Big Thicket National
Preserve, longleaf pine-black gum savannah, 1 0 Sep 1 981 , A. Pecotte 32 (TAMU); NW corner of Big Thicket Bogs & Pineylands
Preserve (TNC), 1 00 ft S of CR 4770, ca. 0.4-0.5 mi E of its jet. with US 69/287 near Lake Hyatt, frequent in moist to wet sandy
loam in portion of boggy wetland with Sphagnum and Sarracenia, 25 Sep 1997, W.R. Carr 17084 (TEX). Wood Co.: in a bog
li N of Mineola, Rte. 37, bog, 29 Sep 1 967, D.5. Correll 35023 (LL). County unknown: damp

flat pinebarrens, 1

)3,C.N.S

ACKNOWLEDGMENTS

I would like to thank the curators and collection managers of the herbaria listed above for providing speci-
mens for this study. I would also like to thank Brenda Molano-Flores for accompanying me on field visits and
Dave Moore of the Kisatchie National Forest for providing information regarding populations in Louisiana.
Thanks also to Geoff Levin, Stephen Downie, Rhiannon Perry, Jenny Cordes, Clark Danderson, Ron Hartman,
and one anonymous reviewer for helpful comments regarding this manuscript. Lastly, thanks to Diane
Szafoni for creating the range map of P. texense and Paul Marcum and Dave Ketzner for testing the key to
Ptilimnium.

REFERENCES

Alvarez, I. and J.F. Wendel. 2(X)3. Ribosomal ffS sequences and plant phylogenetic inference. Molec. Phylogenet.
Evol. 29:41 7-434.

Baldwin, B.G., MJ. Sanderson, J.M. Porter, M.F. Wokiechowski, C.S. Campbell, and M.J. Donoghue. 1 995. The ITS region of

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nuclear ribosomal DNA: a valuable source of evidence on angiosperm phylogeny. Ann. Missouri Bot. Card.
82:247-277.

CoRRELL, D.S. AND M.C JoHNSTON, 1970. Manual of the vascular flora of Texas. Texas Research Foundation, Renner.

CoRRELL, D.S. AND H.B, CoRRELL. 1 972. Aquatlc and wetland plants of the southwestern United States. Environmental
Protection Agency, Washington D.C.

Diggs, G.M., B.L Lipscomb, and RJ. O'Kennon. 1 999. Illustrated flora of north central Texas. Sida, Bot. Misc. 1 6:1 -1 626.

Diggs, G.M., B.L. Lipscomb, M.D. Reed, and R.J. O'Kennon. 2006. Illustrated flora of east Texas. Sida, Bot. Misc. 26:1 -1 594.

Coulter, J.M. and J.N. Rose. 1 909. Supplement to the monograph of the North American Umbelliferae. Contr. U.S.
Natl. Herb. 12:441-451.

Easterly, N.W. 1957. A morphological study of Ptilimnium. Brittonia 9:136-145.

Feist, M.A. and S.R. Downie. 2008. A phylogenetic study of Oxypolis and Ptilimnium (Apiaceae) based on nuclear
rDNA ITS sequences. Syst. Bot. 33:447-458.

Lundell,C.L. 1961. Flora ofTexas. Texas Research Foundation, Renner.

MacRoberts, M.H. and B.R. MacRoberts. 2003. West Gulf Coastal Plain ecoregions. Sida 20:1247-1276.

MacRoberts, M.H., B.R. MacRoberts, B.A. Sorrie, and R.E. Evans. 2002. Endemism in the West Gulf Coastal Plain: im-
portance of xeric habitats. Sida 20:767-280.

Mathias, M.E. and L. Constance. 1 945. Umbelliferae, N. Amer. FI. 288:43-295.

Mathias, M.E. and L. Constance. 1 961 . Umbelliferae. In: CL Lundell, ed. Flora ofTexas, vol. 3. Texas Research Foun-
dation, Renner. Pp. 263-329.

USDA, NRCS. 2010. The PLANTS Database (httpWplants.usda.gov, 1 April 2010). National Plant Data Center,
Baton Rouge, LA 70874-4490, USA.

Weakley, A.S. and G.L Nesom, 2004. A new species of Ptilimnium (Apiaceae) from the Atlantic coast. Sida 21:
743-752.

652

BOOK NOTICE

Dennis W Woodiand. 2009. Contemorary Plant Systematics, 4th Edition. (ISBN 978-1-883-92564-2,
hbk.). Andrews University Press, Sutherland House, 8360 W Campus Circle Drive, Berrien Springs,
Michigan 49104-1700, U.S.A. (Orders: aupo@andrews.edu, 1-269-471-6134, 1-269-471-6224 fax).
$97.99, 666 pp„ color and b&w, 8%" x lUA". Two CDs: University of Wisconsin Photo Atlas of the
Vascular Plants, 3rd Ed. and Interactive Keys to Vascular Plant Families of the World.

Appendix III: Classification of the Flowering Plants as Proposed by the Angiosperm Phylogeny Group (APG)

A NEW VARIETY OF BROMUS FLEXUOSUS
(POACEAE; POOIDEAE: BROMEAE: SECT. BROMOPSIS)
Ana Marfa Planchuelo

Centro de Relevamientoy Evaluacidn de Recursos Agricolas y Naturales
Facultad de Ciencias Agropecuarias. Universidad Nacional de Cdrdoba
CC 509, Cdrdoba (5000), ARGENTINA

RESUMEN

In 1983, Planchuelo described Bromus Jlexuosus as an endemic species for northwestern Argentina. The spe-
cies seemed unrelated to any other species of section Bromopsis Dumort. cited for the country in the available
floras (Burkart 1969; Camara Hernandez 1970, 1978). Along with the original description, Planchuelo (1983)
separated the specimens into two altitudinal ecotypes, which were identified as Group I and II, leaving the
taxonomic status of each Group for future investigation. In the following treatments of Bromus for Argen-
tina (Zuloaga et al. 1994; Gutierrez & Pensiero 1998) and for the Americas (Planchuelo & Peterson 2000;
Pavlick et al. 2003), no infraspecific taxa were considered for B. Jlexuosus. More recently, during a revision
of Bromus toward a preparation of a monograph for South America and with the aim to provide a better
understanding of the species of section Bromopsis growing in the northwestern Argentina and the Central
Andes, current literature and specimens of different herbaria were reviewed. Additional collections of B.
Jlexuosus were reported for Peru (Saarela et al. 2006) and for Bolivia (Planchuelo 2010) which expanded the
area of distribution of the species northward in the Andes. The affinities (panicles and pedicels flexuosity)
and the differences (sizes and pubescence of glumes and lemmas) between B. Jlexuosus and other species of
section Bromopsis were defined in a partial review of species for the Central Andes (Planchuelo 2009).

all having a flexuous panicle, a phenetic cluster analysis based on morphological characters, as well as an
evaluation of discriminant characters was performed on herbarium specimens.

Based on the cluster analysis and selected discriminant characters, I present evidence for two variet-
ies. A detailed description, habitat information, geographical distribution, representative specimens, and
illustration for B. Jlexuosus var. Jamatinensis are provided. Also, taxonomic keys to identify the varieties of
B. Jlexuosus and the related taxa are included.

J. Bot Res. Inst Texas 4(2): 653 -662. 2

MATERIALS .

METHODS

The morphological data were obtained from herbarium specimens as cited. Vegetative and floral structures
were included in the data matrices in order to establish the relationship between the studied materials and
the taxa that they represent. The taxa considered were B.Jlexuosus, B. lamtus, and B. modestus. References for
identification are Hitchcock (1927), Planchuelo (1983), Pinto-Escobar (1981, 1986), Matthei (1986), Tovar
(1993), Renvoize (1994, 1998), Gutierrez and Pensiero (1998), and Planchuelo and Peterson (2000). Bromus
pitensis considered by Planchuelo (1983) as a related species of B.Jlexuosus was not considered in this study
because it does not have the typical flexuous pedicels (Planchuelo 2010) and the nodding panicles make
the species more related to B. segetum Kunth.

Measurements for each variable were taken with the use of 10-15x magnifying lens, and the more
frequent value (mode) of that character for each specimen was recorded. Measurements of first and second
glumes were recorded independently; however, pubescence of both glumes were considered as only one

distribution on main culm was coded as: 1 = leaves only at the base of the plant; 2 = leaves at the base of
the plant and along the culms. Leaf blade pubescence for each surface was divided into three states from
1 to 3: 1 = glabrous; 2 = scattered pubescent along nerves and margins; 3 = scattered to densely pubescent
on all surfaces. Pedicel pubescence was divided into three states: 1 = glabrous; 2 = scabrous to scattered
pubescent; 3 = densely pubescent to lanate. Glumes pubescence was divided into three states: 1 = glabrous to
glabrate; 2 = evenly scattered pubescent; 3 = pubescent all over and dense at the base and margins. Lemma
pubescence was scored for three states: 1 = glabrous to glabrate; 2 = scattered pubescent and villous only at
the base; 3 = dense pubescent all over and villous at the base and along margins. Maximum and minimum
values, as well as mean (p) and coefficient of variation (CV), expressed in percentage were calculated for all
continuous variables. The mode (M) was calculated for discrete variables such, leaf distribution, number of
spikelets, florets, and nerves.

The results presented here are based on the analysis of a matrix containing the following 17 morpho-
metric parameters: plant height; leaves disposition in the plant; blade width and pubescence above and
below; panicle length and number of spikelets; pedicel pubescence; spikelets length and number of florets;
first glumes length, second glumes length; pubescence of glumes, second lemma length, number of nerves’
pubescence and awn length. The cluster analysis (Sneath & Sokal 1973) was base on 39 Operational Taxo-
nomic Units (OTU) using Manhattan distance coefficient and the unweighted pair grouping method of
arithmetic average (UPGMA) (Michener & Sokal 1957).

Specimens included m the multivariate analysis are identified with the OUT’s identification number
between brackets (e.g., [10]). Data on geographical distribution, elevation and habitat are based on herbarium
specimen labels and personal field observations.

RESULTS /

J DISCUSSION

Most of the characters have low (<20% CV) to moderate (<40% CV) variability; except the character “plant
height- which CV was over 40% (Table 1). Small plants and narrow blades of B.^Iexnosas are usually associ-
ated With specimens collected at high altitude and they are correlated with species habitat as it was reflected
by the relatively low CV (less than 20%) found when the statistical analysis was done with selected samples
trom similar altitudinal range. Panicle length shows very good correlations with plant heights; however, its
CV is less than 30% in all three species. The comparison of panicle length, number of spikelets and plant
height shows that Bromus lanatus and B. modestus have lower number of spikelets and much larger plants
than B.Jlexuosus given the same length of panicles. These relationship of characters along with large sizes of

the main discriminant components that separate the three species in two clusters (Fi^lJ^BlmrsTode^^^^
is the only species with leaves present only at the base of the plant; therefore, this feature along with the
lack of pubescence in pedicels and glumes, are the main discriminant characters between this species and

656

1.5 1.0 0.5 0.0

B. lanatus. The differences between varieties of B.flexuosus are glumes sizes and pubescence distribution in
blades and glumes.

The results shows that, although the taxa have some overlapping in vegetative and floral measurement
and in some cases they have similar pubescence characteristics (Table 2), the combination of all the features
in the cluster analysis (Fig. 1) clearly separates the 39 analyzed specimens into two main groups. One cluster
comprises all representative OTUs of B.flexuosus divided into two secondary branches that represent the two
varieties; and the other main cluster has two secondary branches that correspond to the specimens identified
as B. lanatus and B. modestus. It is important to point out that in this review the specimen Kurtz 1672a (CORD)
cited as B.flexuosus in Planchuelo (1983) are now cited as B. modestus, species which was not described at the
time of the publication of B.flexuosus. This new identification approach allowed a better taxa definition, as it is
presented in the following taxonomic treatment.

TAXONOMIC TREATMENT

KEY TO THE SPECIES BROMUS FLEXUOSUS, B. LANATUS, AND B. MODESTUS
1 . Panicles (8-)1 0-20(-25) cm with (8-) 1 0-25 spikelets and (5-)6-8 florets; first glumes (7-)8- 1 0 (1 1 ) mm long,
second glumes (8-)9-11(-12) mm long; lemmas 11-14 mm long, 7-nerved, the awns (4.5-)6-7(-8) mm

long __1. B. flexuosus

1. Panicles (3-)4-8(-1 1) cm long with (3-)4-8(-10) spikelets and 3-5(-6) florets; first glumes 5-7 mm long,
second glumes 7-8.5 mm long; lemmas 7-10(-11) mm long, 5-nerved, the awn 2-3.50 mm long.

2. Plants with basal and cauline leaves; pedicels lanate 2. B. lanatus

2. Plants with basal leaves only; pedicels glabrous to scabrous, less

; frequent scattered pubescent 3. B. modestus

657

Table 2. Comparison of Attribute Variables indicating: the number of specimens which have the coded charaaer, from the total number of specimens used in the discriminant
analysis.

Bromus lanatus n = \2

2 4 2

1. Bromus flexuosus Planchuelo, Kurtziana 16:123-131. 1983.

Caespitose perennials. Culms (15-)30-60(-70) cm tall, base covered with loose non fibrous sheaths and
leaves. Leaf sheaths retrorse pubescent or scabrous; ligules 1. 5-2.5 mm long, lacinate; auricles 1-2 mm long,
caducous; blades (10-)20— 30(-50) cm long x (1.5-)3.5-6(-7) mm, glabrous, scattered pubescent in nerves
to densely pubescent above and below. Panicles (8-)10-20(-25) cm long, open, with (8-)10-25 spikelets,
branches and pedicels very flexuous, scabrous, scattered pubescent to lanate. Spikelets (16-)18-20(-24) mm
long, with (5-)6-8 florets, slightly imbricate, internodes visible at maturity; glumes narrow-triangular; first
glumes (7-)8-10(-ll) mm long, 1-nerved; second glumes (8-)9-ll(-12) mm long, 3-nerved, mucronate,
scattered to pubescent all over and more dense at the base and margins; lemmas 11-14 mm long, ovate,

mm long. Paleas almost equal to the lemma in length; anthers 0.8-1. 2 mm long. Caryopsis 10-11 mm long,
widely furrowed not adhering to the palea or lemma.

The two proposed varieties can be easily identified by the following key:

KEY TO THE VARIETIES OF BROMUS FLEXUOSUS

1 . Blades glabrous or sometimes sparsely pubescent along the nerves and margins above, sparsely to densely pubescent
below; panicles 8-13(-18) cm long; glumes evenly scattered pubescent, the first (8-)9-I0(-n)mm long, the second

1 . Blades scattered to densely pubescent ab
below; panicles (1 3-)1 5-25 cm long; glun
7-8(-9) mm long, the second 8-1 0(-1 1) r

e, glabrous or sparsely pubescent along the nerves and margins
s pubescent all over and dense at the base and margins, the first

n B. flexuosus var flexuosus

r. famatinensis Planchuelo, var. nov. (Fig. 2B).
La Vega de la Mesada, (28°58'S, 67°47'W) 3650, m 15-16 Mar

i. (15-)20-40(-50) Cl

Etymology.— The epithet “famatinensis” was chosen to reflect the major region of distribution, and specially
the place where this new variety grows.

DistrihutionandHohitat.— NativeinthemountainsofnorthwestemArgentinaandinthecentralAndesofBohvia

and Peru. The new variety grows in the same mountain slopes as the typical variety but at altitudes above 3000 m.

Comments.— The specimens Burkart & Troncoso 11920 and Parodi 7976 were erroneously cited by Gutier-
rez and Pensiero (1998) as B. lanatus. The specimen Kurtz 1672a (CORD) cited as B. flexuosus in Planchuelo
(1983) are now cited as B. modestus.

658

Planchuelo.Ar

ia el Cerro Manchado,

lurkart & Troncoso 11920 (BAA). L
>arodi 7976 (BAA, US); En las cercanias de la mina El Oro, 23/25 Jan 1879 Hieronymus 433 (CORD[17]). Famatina: La vega de La Mesada,
a. 3630 m, 15-16 Mar 1906, Kurtz 13894 (CORD[12|, US); Mina San Juan, 3050-3200 m, 21 Feb 1906 Kurtz 13602 (CORD|15l); Bajada
ntre la Cumbre de la Cuesta de La Mesada y La Cienaga de los Arenales, 3600-3750 m, 21 Mar 1906, Kurtz 13989 (CORD); Cienaga
le la Calera, 3600-3650 m, 20 Mar 1906, Kurtz 13937 (CORD[14)); La Mesada, rio Amarillo superior. 3500, 25 Mar 1906, Kurtz 13836
y 13836 bis (CORD); Las Trancas, Cerro Coloradito, 12 Mar 1907. leg. R Tejada s/n, Kurtz 14595 (CORDH61); Real Viejo, 6 Mar 1907,
Curtz 14758 (CORD); Cueva de P6rez, 26/28 Jan 1879, Hieronymus et Niederlein 401 (CORD[18|). Dpto. General Sarmiemo, El Volc4n, 28
50 (CORD1131); Las Cortaderas, entre El PeflOn y El JagCiel. 27 Feb
e, Cerca camino a San Miguel, quebrada Sikipampa, 4000 m, 2 Jan

Distribution and Habitat. — Native in Northwestern Argentina and in the Central Andes i
Peru. It grows in grassland and mountain slopes above 2000 m.

Major references . — Planchuelo (1983); Zuloaga et al. (1994); Gutierrez and Pensiero (1998); Planchuelo
and Peterson (2000); Pavlick et al (2003).

Comments. — ^The specimen Peterson & Refulio-Rodriguez 16556 (ACOR, US, USM) was cited for Peru in
Saarela et al. (2006) and was the first record of B.flexuosus beyond noi
nally described.

Veg. 2:708. 1817. Type: ECUADOR. Pichincha/N>

BM; photo K!. US-865493 fragm!).

Bromus oUganthus Pilg., Bot. Jahrb. Syst. 25:718. 1898. Type Protologue: Ecuador. Imbabura: Paramo
(l^CTOTYPE, designated here: US-00131962!; bouctotype: US-0008161 11), ECUADOR. In paramos :
de la Media Luna, alt. 4400 m, Stubel 230a (Syntype: B), ECUADOR. Pichincha. Sfuhel 20c (syntype: 1
Puntas, alt. 4400 m. Stubel 207a (syntype: B). COLOMBIA. Crescit in monte Tolima ad Boca del K

Perennial. Culms (10-)15-20(-30) cm tall. Leaf sheaths close and retrorsus pubescent to the ape:

Journal of the Botanical Research Institute of Texas 4(2)

membranous glabrous, apex dentate; blades 4-8 cm x 1-3.5 mm, scattered to densely pubescent above and
below with long hairs at the margins. Panicles (4-)6-8(-ll) cm long, with (4-)5-8(-10) spikelets, branches
flexuosus shorter towards the apex, pedicels lanate and flexuosus. Spikelets 13-16 mm long, 4-5(-6)-florets
not imbricates with rachilla visible; glumes lance-attenuate, pubescent throughout, densely so near base and
margins, hrst glumes 5-7 mm long, 1-nerved, second glumes Z.5-8.5 mm long, 3-nerved; lemmas 9-11
mm long, oval-lanceolate, 5-nerved, densely pubescent and villous at the base and margins, awns 3-3.5
mm long, subapical. Paleas a little shorter that the lemma; anthers 0.5-0.8 mm long. Caryopsis 6 mm long,
widely furrowed, not attached to the lemma and palea.

Distribution and Habitat.— Native to the central Andes. Commonly occurs in grass paramos (treeless
alpine plateau, neo-tropical ecosystems) on sandy to rocky slopes and humid places above 2000 m; found
in northern Argentina and Chile and especially in Bolivia, Peru, and Ecuador.

Major references.— Standky (1947), Pinto-Escobar (1981, 1986), Matthei (1986), Tovar (1993), Gutierrez
and Pensiero (1998), Renvoize (1998), Planchuelo and Peterson (2000).

Comments.— 1 agree with the original description of Kunth (1816) and with Matthei (1986) that B.
lanatus has lemmas with hve nerves. The observation “lemma 7-nervada” from dry fragments (isotype) of
B. lanatus by Planchuelo (1983) is erroneous because what was thought to be tbe two laterals emerging and
inconspicuous nerves were the marked crest of the folded lemma base, revealed only after hydrating the
materials. The specimen Spegazzini 2558 from the herbarium BAA has five plants and two panicles mounted
in the same sheet. The plant mounted at the left bottom corner, assigned to the letter A is B. lanatus, all
other plants and panicles correspond to B. flexuosus var. flexuosus. This important finding indicates that
both species share the same habitat in Salta, Argentina. Unfortunately, that region is not well explored and
no new specimens have been collected. A similar case of both species sharing the same habitat is seen in
three collections of the BAA herbarium by Parodi in “La Rioja, Famatina, Camino a La Mexicana” where the
specimen numbered Parodi 7919 is B. lanatus, the specimen Parodi 7963 is B. flexuosus var. flexuosus and Parodi
7976 is B. flexuosus var. famatinensis.

661

3800 m. Rageland, Jalca areas of saturated soil, 17 Feb 1983, Smith & Vasquez 3483 (MO[29l); 52 km N of Cajamarca on HWY 3, N towards
Bambamarca, 3700 m, 16 Mar 2000, Peterson & Refulio- Rodriguez 14908 (MO). Cuzco: Paucartambo, 21 km NE of Paucartambo on road
to Ires Cruces, 3460 m, 13°ir54.8"S. 71°38'40.5"W, 18 Mar 2002, Peterson & Refulio-Rodriguez 16622 (ACOR [20|, US). Tacna: 7 km
NW of Alto Peru on Rio Uchusuma, 4330 m, 13 Mar 1999, Peterson, Refulio Rodriguez& Salvador Perez 14750 (ACOR[211, US).

Perennial caespitose with the base covered with loose non fibrous sheaths. Culms 10-35 cm tall, with leaves
only at the base of the plant. Sheaths often open at middle way, retrorse pilose to sericeus, ligules membra-
nous, glabrous, 1 mm long, apex laciniate. Auricles absent. Blades 3.5-11 cm long x 1-3 mm folder or plane,
glabrous, glabrate or scattered to densely pubescent along nerves and margins on both sides. Panicles lax,
3-8 cm long, branches short and flexuosus, with 3-6 spikelets, pedicels flexuosus, glabrous to scabrous,
less frequent scattered pubescent. Spikelets 10-15 mm long 3-4(-6)-florets, imbricate. Glumes glabrous,
glabrate to scattered pubescent, the first narrowly-triangular 5-7 mm long, 1-nerved, the second lanceolate
7-8 mm long, 3-nerved. Lemma lanceolate, 7-9 mm long, 5-nerved, glabrous or glabrate to scattered pu-
bescent all over and villous only at the base, apex acute; awn subapical, straight 2-3.5 mm long; Palea 6-8
mm long with long hairs on the nerves. Anthers 1 mm long. Caryopsis 6-8 mm long narrowly furrowed
attached to the lemma and palea.

Distribution and Habitat. — Native in the Andes, from northern Chile and Argentina northward to Ecuador
and Colombia. Grows on mountain slopes and high plains, between 3600-4700 m.

Major references. — Renvoize (1994, 1998); Planchuelo & Peterson (2000).

Comments. — Renvoize (1994) described B. modestus by giving a new name to B. frigidus Ball. Some
authors (Pinto Escobar 1981 and Tovar 1993) considered B. frigidus as synonym of B. pitensis. Saarela et al.
(2007) demonstrated that B. modestus and B. lanatus formed an exclusive clade in a plastid tree, based on
chloroplast and nuclear DNA sequence data, which reaffirms the affinities of both species established by the
morphological analysis in this paper. The specimens, Weberbauer 6902 (US) and leg. Harlan, US 727045 (US),
were cited as B. lanatus by Standley (1947), the specimen Parodi 10854 (BAA, US) was also cited as B. lanatus
by Gutierrez & Pensiero (1998) and the specimen Kurtz 1672a (CORD) was cited as B. flexuosus in Planchuelo
(1983).

ACKNOWLEDGMENTS

The author is indebted to the curators of the herbaria cited for the loan of several specimens, and for their kind
attention during my visit to their institutions. I thank Victoria Hollowell for her valuable suggestions, to Paul
Peterson and the anonymous reviewer for their comments and suggestions that have greatly improved this
paper. This research was partially funded by SECYT of the Universidad Nacional de Cordoba, Argentina.

REFERENCES

Burkart, a. 1 969. Grami'neas. Flora ilustrada de Entre Rios (Argentina). Institute Nacional deTecnologta Agropec-
uaria, Buenos Aires 6(2):1-551.

Camara Hernandez, J. 1970. Bromus. In: A.L Cabrera. Gramineas. Flora de la Provincia de Buenos Aires. Institute
Nacional deTecnologia Agropecuaria, Buenos Aires 4(2):85-101.

Camara Hernandez, J. 1978. Bromus. In: E.G. Nicora. Gramineae. Flora Patagonica. INTA, Buenos Aires 3:77-93.
Gutierrez, H.F. and J.F. Pensiero. 1 998. Sinopsis de las especies Argentinas del genero Bromus (Poaceae). Darwiniana
35:75-114.

Hitchcock, A.S. 1 927. The grasses of Ecuador, Peru, and Bolivia. Contr. U.S. Natl. Herb. 24(8):291 -556.

Kunth, CS. 1816. Bromus. In: A. Humboldt, A. Bonpland, and CS. Kunth. Nova genera et species plantarum 1.

Facsim. ed. Weinheim (J. Cramer) 1 963. Pp. 1 50-153.

Matthei, 0. 1986. El genero Bromus L. (Poaceae) en Chile. Gayana Bot. 43:47-1 10.

MicHENEa C.D. and R.R. Sokal 1 957. A quantitative approach to a problem of classification. Evolution 1 1 :490-499.
Pavlick, L.E., A.M. Planchuelo, P.M. Peterson, and R.J. Soreng. 2003. Bromus. In: RJ. Soreng, P.M. Peterson, G. Davidse,

IV. Subfamily Pooideae. Contr. U.S. Natl. Herb. 48:1 54-1 91 .

PiNTO-EscoBAa P. 1981. The genus Bromus in northern South America. Bot. Jahrb. Syst. 102:445-457.
PiNTo-EscoBAa P. 1 986. El genero Bromus en los Andes Centrales de Suramerica. Caldasia 1 5(71-75):! 5-34.
Planchuelo, A.M. 1983. Una nueva especie de Bromus (Poaceae) de Argentina. Kurtziana 16:123-131.

Planchuelo, A.M. 1 991 . Estudios sobre el complejo Bromus catharticus (Poaceae), I. Evaluacion estadfstica de los
caracteres taxonomicos. Kurtziana 21 :243-257.

Planchuelo, A.M. 2009. Nueva distribucion de especies de Bromus (Poaceae: Bromeae) p
Andina. Bol. Soc. Argent. Bot. (Supl.) 44(3-4):204-205.

Planchuelo, A.M. 2010. Nuevas citas de especies de Bromus L. (Poaceae: Bromeae) para Bolivia. Kempffiana
6(1):3-15.

Planchuelo, A.M. and P.M. Peterson. 2000. The species of Bromus (Poaceae: Bromeae) in South America. In: S.W.L
Jacobs and J. Everett, eds. Grasses: Systematics and Evolution. CSIRO, Collingwood, Australia, Pp. 89-101 .
Renvoize, S.A. 1 994. Notes on Sporobolus & Bromus (Gramineae) from the Andes. Kew Bull. 49:543-546.

Renvoize, S.A. 1998. Gramineas de Bolivia. The Royal Botanic Gardens, Kew, U.K.

Saarela, J.M., P.M. Peterson, and N.F. Refulio-Rodriguez. 2006. Bromus ayacuchensis (Poaceae: Poideae: Bromeae), a
new species from Peril, with a key to Bromus in Peru. Sida 22:91 5-926.

Saarela, J.M., P.M. Peterson, R.M. Keane, J. Cayouette, and S. W. Graham. 2007. Molecular phylogenetics of Bromus
(Poaceae: Pooideae) based on chloroplast and nuclear DNA sequence data. In: Columbus, J.T., E.A. Friar, J.M.
Porter, LM. Prince, and M.G. Simpson, eds. Monocots: comparative biology and evolution-Poales. Rancho
Santa Ana Botanic Garden, Claremont, California. Pp. 450-467.

Sneath, P.H.A. and R.R. Sokal. 1973. Numerical taxonomy: the principles and practice of numerical classification.
San Francisco: Freeman.

Standley, P.C. 1947.Gramineae. Grass family. In: Mac Bride, Flora of Peril, Part. I. Field Mus Nat Hist Bot Ser
13:115.

TovAa 6. 1 993. Las Gramineas (Poaceae) del Peru. Ruizia 1 3:1 -480.

Zuloaga, F.O., E.G. Ncora, Z.E. Rugolo de Agrasai^ O. Morrone, J. Pensiero, and A.M. Cialdella. 1 994. Cat^logo de la familia
Poaceae en la Repilblica Argentina. Monogr. Syst. Bot. Missouri Bot. Gard. 47:i-xi, 1-1 78.

THE GENUS RYTIDOSPERMA (POACEAE)

Stephen J. Darbyshire

IN THE UNITED STATES OF AMERICA
Henry E. Connor

1939; Darbyshire & Connor 2003; Dean 6
The genus Rytidosperma is distinguis
transverse rows of hairs on the back of th<

nly in California, Oregon and Hawaih (Whitney e

3m Danthonia sensu stricto primarily by the presence of
a (Figs. 1-3). These hairs may be in discrete tufts or form
more or less continuous lines. One row occurs towards the base just above the callus and the other towards
the apex just below the awn sinus. Rows, especially the upper, may be reduced to tufts only at the margins.
Other hairs may or may not be scattered across the back of the lemma between the rows. In some species the
lemma back indumentum may be lacking altogether, while in other species the lemma back is more or less
evenly covered by hairs with no distinct trace of the two rows of tufts. Ensheathed spikelets, cleistogenes,
were reported by Chase (1918) in the leaf axils of many species of Danthonia (including “D. semiannularisl,
but these are absent from Rytidosperma (Vickery 1956; Connor & Edgar 1979). Florets of the exserted inflo-
rescences may be either chasmogamous or cleistogamous (Edgar & Connor 2000).

Descriptions and measurements given here follow the conventions of Connor & Edgar (1979) and
are based primarily on plants of North American and Hawaiian origin. Meiotic chromosome counts of R.
racemosum and R. richardsonii were determined on plants grown from seed obtained from collections origi-
nating in California. These counts are indicated in square brackets after the original collection citation.
Voucher specimens for chromosome counts of R. racemosum and R. richardsonii are deposited at DAO with
associated material at DAO, CHR and UC. Counts of plants of New Zealand provenance are presented for
R. caespitosum; vouchers are deposited at AK and CHR. Illustrations of lemmas are prepared from North
American (line drawings) and New Zealand (color paintings) material.

This study is primarily focused on species that have become naturalized in the United States; however,

ell have occurred.

icluded since undetected e;

i. Culms (1.5-)30-90(-140)

provide examples of poorly known species, or when they provide information on introduction sit
TAXONOMIC TREATMENT

Rytidosperma Steud., Syn. PI. Glum. 1:425. 1854. Type: Rytidosperma lechleri Steud.

l^oiodanthoma Zotov, New Zealand J. Bot. 1:104. 1963. Type: Notodanthonia unarede (Raoul) Zotov.

Austrvdanthorua H.E Under, Telopea 7:269. 1997. Type AustrodaKthoma caespnosa (Gaudich.) H.P Under.

Plants perennial, densely or loosely cespitose, sometii
cm, erect or nodding. Leaves usually mostly basal; s
apices usually with tufts of hairs, sometimes extending across'.he collar; ligules a rto orhahsrblaies
persistent or disarticulating, flat, involute or convolute, glabrous or variously pubescent. Inflorescences
terminal, racemes or panicles. Spikelets somewhat laterally compressed, with 3-10 florets- florets bisexual
chasmogamous or cleistogamous, terminal florets reduced; disarticulalion above the glumes and between
the florets; glumes (2-)8-20 mm long, subequal or equal, usually exceeding the florets (excluding awns),
sti fly membranous, 3-13 veins, usually with scarious margins; calluses sharp or somewhat blunt with
latera tufts of stiff hairs, disarticulation oblique; lemmas (the main lemma body) ovate to lanceolate with 2
complete orincomplete transverse rows of tufts of stiff hairs, sometimes reduced to marginal tufts 5-9(-ll)
veins, apices bilobed, the lobes usually at least as long as the body, acute or acuminate with a long awn-
hke arista; a central awn between the lobes and longer than them, usually proximally twisted below into a
column, usually geniculate or reflexed; lodicules 2, fleshy, usually with apical hairs or glabrous; anthers
vary greatly in size depending on whether they are from cleistogamic or chasmogamic flowers (usually <
1.2 mm and > 1.8 mm, respectively). Caryopses 1.2-3 mm long, obovate to elliptic, golden to dark brown,
ree, hila punctiform to somewhat elliptic. Cleistogenes not formed. x2 = 12 (Murray et al. 2005) Name
from the Greek rhytidos, “wrinkles,” and sperma, “seed” (vide Connor & Edgar 1979).

Darbyshire et al., The genus Rytidosperma in the U.S.A.

665

KEY TO THE SPECIES OF RYTIDOSPERMA NATURALIZED IN
. Upper lemma hairs in isolated tufts or at margins only and not forming a
2. Callus of second lemma 0.5-1.

2. Callus of second lemma 1-1.5 r

E UNITED STATES OF AMERICA

1 long, its hairs usually overlapping the lower ro\

gradually narrowed to a fine awn-like arista

)ng, its hairs rarely or just reaching lower row of le

lemma lobes 5-10 mm long, abruptly narrowed to a fine avyn-like arista

. Lemma hairs in two continuous transverse rows of tufts, with or without hairs between tt

3. Central awn 10-20 mm long, column 4-5 mm long

3. Central awn usually less than 10 mm long, column 0.5-3 mm long.

Timas 1.8-2.4mmlong;av
jally not reaching upper rc
Timas 3-4 mm long; awn
?riapping upper row

n 2.5-3 I

1 long, sparingly twisted; Ic

Rytidosperma biannulare (Zotov) Connor & Edgar, New Zealand J. Bot. 17:324. 1979. (Fig. 1C, 3B). Noto-

Plants caespitose. Culms 30-85 cm, erect, smooth and glabrous, glabrous below inflorescence, branching
intravaginal. Leaves mostly basal, exceeded by the culms, flag leaf blades usually reaching or exceeding the
inflorescences; sheaths mostly glabrous, often purplish distally, apical tufts of hairs to 5 mm or sometimes
absent; ligules 0.3-0.5(-l) mm; blades 30-40 cm long, to 5 mm wide, usually involute, margins, apices.

Inflorescences paniculate, 10-20 cm long, narrow and compact; rachis scabrous; pedicels shorter than
spikelets, scabrous. Spikelets (7-)10-15 mm long, with 6-7 florets; glumes 7.5-ll(-13.2) mm long, surpass-
ing florets, subequal, lanceolate, acute, glabrous, light green to stramineous usually purple at margins and
apex; lower glumes 5-7(-9) veins; upper glumes 5 veins; rachilla segments 0.3-0.5 mm long; calluses
0.5-0.7 mm long, hairs to about 1 mm and reaching the lower lemma hairs; lemmas 1.8-2.4(-2.8) mm long,
hairs of lower row usually not or only just reaching the upper row, lower row sometimes ill-defined, hairs
of upper rows reaching or surpassing the awn column apex but not the apex of the lemma lobes, with short
scattered hairs (rarely glabrous) between the rows; lobes 3.5-5(-8.5) mm long, acuminate; awn 6-10(-12.5)
mm long, twisted column 2.5-3 mm long; paleas 2.5-4.6 mm long, exceeding the lemma sinuses, obovate,
emarginate, sparsely hairy between the veins, margins usually with long hairs, veins ciliate; anthers 0.8-1 .6
mm long. Caryopses 1. 2-1.9 mm x 0. 6-0.8 mm; embryos 0.5-0.8 mm long; hila 0.3-0.6 mm long. 2n =
48; New Zealand plants (Murray et al. 2005).

A single collection from southwestern Oregon (Peck 23954) and another from Maui (Hobdy 2389) provide
the only evidence that this species may have naturalized in the United States, although cultivated specimens
from Santa Cruz Co., California, have also been seen (Fig. 4A). The species has been grown experimentally
in North America under the name Danthonia semiamularis (Labill.) R. Br., possibly as early as 1905 (Wein-
traub 1953). However, as pointed out by Vickery (1956), D. semiannularis is a name that “has been used for
almost any but the true species” and more than one species was probably imported to North America under
this name. Since tetraploid R. biannulare is regarded as native to New Zealand and unknown in Australia,
this species may have been more recently introduced than the early 20th Century importation of Australian
species.

Whitney et al. (1939) and Wagner et al. (1999), respectively, reported R. semiannulare as introduced to
the state of Hawai'i (on Molokai) in 1903 and first collected (on Maui) in 1937. The 1937 specimen (Hosaka
1767) is, however, referred to R. caespitosum. Label data on a relatively recent collection of R. biannulare from
West Maui (Hobdy 2389) suggest that this species has naturalized there.

Specimens examined: CALIFORNIA. Santa Cruz Co.: Ives plots near Aptos, Corralitos area, Icultivated], 20 May 1940, P.B. Dicfeey 920.

Journal of the Botanical Research Institute of Texas 4(2)

Darbyshire et al., The genus Rytidosperma in the U.S.A.

667

D920 (AHUC, US). HAWAI‘1. Maui: West Maui, Hanaulaiki, elev. 3500 ft, May 1985, R. Hobdy 2389 (BISH). OREGON. Curry Co.: 5
mi S of Gold Beach, shady bank, 23 Jul 1945, M.E. Peck 23954 (WILLU).

Rytidosperma caespitosum (Gaudich.) Connor & Edgar, New Zealand J. Bot. 17:325. 1979. (Fig. 2B, 2C,

Plants densely to loosely caespitose, sometimes shortly rhizomatous. Culms 43-80 cm, erect, smooth and
glabrous, glabrous or sparsely scabrous immediately below the inflorescence, branching intravaginal (or rarely
extravaginal). Leaves mostly basal, usually exceeded by culms, flag leaves sometimes reaching inflorescence;
sheaths glabrous or pilose, apical tufts of hairs 1-4 mm long, sometimes scanty; ligules (0.3-)0.5-1.2 mm
long; blades 6-25 cm long, 1.5-3 mm wide, involute or more or less flat, glabrous or variously pubescent.
Inflorescence paniculate, 5-1 1 cm long, linear to ovate, more or less compact; rachis scabrous to somewhat
pubescent; pedicels shorter than spikelets, scabrous to somewhat pubescent. Spikelets 10-20 mm long,
4-9 florets; glumes 9-18(-20) mm long, surpassing florets, subequal to unequal, lanceolate, acuminate,
glabrous or sometimes with scattered long hairs, light green to stramineous often purple at margins and apex;
lower glumes (l-)3-5(-7) veins; upper glumes (3-)5-7(-9) veins; rachilla segments 0. 1-0.4 mm long;

about 3.5 mm long and reaching or surpassing the palea apex but not apex of awn column or lemma lobes,
glabrous between the rows; lobes (6-)7-10 mm long, acuminate or aristate; awn 10-20 mm long, tightly
twisted column 4-6 mm long; paleas 2. 5-5.5 mm long, surpassing lemma sinus, lanceolate to obovate,
emarginate, glabrous between the veins, margins with a few long hairs, veins ciliate; anthers 0.5-2. 6 mm
long. Caryopses 1.7-2. 3 mm x 0.8-1. 1; embryos 0.7-1 mm long; hila 0.25-0.7 mm long. 2n = 24, 48, 72;
Australian plants (Abele 1959; Brock & Brown 1961; Waters et al. 2010). 2n = 24; New Zealand plants (B.G.
Murray & J.P. de Lange, hie comm.; AK 25913C, CHR 549710).

Grasslands, pastures, rangelands and disturbed areas up to 200 m. A highly variable and widespread
species with co-occurring polyploid races indigenous to southern Australia (Vickery 1956; Abele 1959;
Brock & Brown 1961; Waters et al. 2010). Rytidosperma caespitosum has naturalized at a few scattered loca-
tions in California (Berkeley, Pescadero and San Diego regions), and has been cultivated at Pullman, WA,
and several sites in California (Fig. 4B). Two collections form Hawai'i by Hosaka in the 1930s are referred
to this species, but the lack of recent collections suggests that it may not have persisted. It has also been
naturalized in New Zealand since at least 1892 (Zotov 1963). It is listed, with some uncertainty, among the
species introduced to the United Kingdom through discarded wool-waste (Lousley 1961; Ryves 1988).

iintheU.S.A.

670

Journal of the Botanical Research Institute of Texas 4(2)

Plants densely to loosely caespitose to somewhat spreading, shortly rhizomatous. Culms (22-)30-90
cm, erect, usually smooth and glabrous, usually scabrous-pubescent immediately below the inflorescence,
branching extravaginal, cataphylls scaly. Leaves mostly basal and greatly exceeded by the culms, flag leaf
blades usually not reaching the inflorescence; sheaths densely hairy or glabrous, apical tufts of hairs 1-3.5
mm long; ligules (0.1)0.3-1 mm long; blades to 30 cm long and to 5 mm wide, flat, folded, or involute,
glabrous, scabrous or pubescent. Inflorescences racemose or paniculate, (3-)4-10 cm long, ovate to lin-
ear, compact; rachis scabrous to finely pubescent; pedicels shorter than the spikelets, glabrous, scabrous
or finely pubescent. Spikelets 9-15(-18) mm long, 5-7(-10) florets; glumes (7,3-)8-13(-17.5) mm long,
surpassing florets, subequal, lanceolate, glabrous, scabrous, or sometimes with scattered hairs, light green
to stramineous often purple at margins and apex; lower glumes 5-9(-ll) veins; upper glumes 5-7(-9)
veins; rachilla segments 0.2-0.5 mm long; calluses 0.5-1.3 mm long, hairs about 1.5 mm long and usually
reaching the lower lemma hairs; lemmas (2-)2.5-4.2 mm long, 9(-ll) veins, lower row of hairs continu-
ous or with weak (rarely absent) central tufts, hairs of the marginal tufts usually reaching the upper row of
hairs, upper row of hairs composed of 2 marginal tufts, sometimes with 2 additional scanty tufts between,
hairs reaching or slightly exceeding the base of the awn, otherwise glabrous; lobes 5-ll(-13) mm long,
acuminate, anstate; awn (7-)9-17.5 mm long, tightly twisted column 1.5-4 mm long, somewhat reflexed
at base and revealing palea apex or not; paleas 3-6 mm long, exceeding the lemma sinuses, emarginate,
intercostal region glabrous or scabrous, margins glabrous or with sparse long hairs, veins ciliate; anthers
0.4-2.7 mm long. Caryopses 1.8-2.5(-3) mm x 0.8-1. K-1.6) mm; embryos 0.7-K-1.5) mm long; hila
(0.3-)0.4-0.5(-0.7) mm long. 2n = 24; Australian plants (Abele 1959; Brock & Brown 1961) and Californian
plants (Myers 1947 [as Danthonia pilosa]).

Commonly growing on dry, nutrient-poor soils. Habitats include pastures, rangelands and disturbed
areas at elevations up to about 800 m in California, and about 1675-2840 m in Hawai'i. It is a common
weed in coastal regions of California and southwestern Oregon (Fig. 4C). At best, only of moderate forage
value in the United States where it is usually considered a troublesome pest competing with more desirable
species (Murphy & Love 1950; Stone et al. 1992).

An Australian species, R. penicillata has been incorrectly known in the United States for many years
under the name Danthonia pilosa R. Br. (e.g., Hitchcock 1951; Sampson et al. 1951; Weintraub 1953). Various
common names have been used, including hairy danthonia, hairy oatgrass, Australian oatgrass and poverty
grass. Although it is considered a poor quality forage grass, it was introduced and tested in the continental
states in 1911 and again in 1921 (Weintraub 1953). By the 1940s it had become a troublesome weed at scat-
tered localities mainly in coastal areas from southwestern Oregon to central California (Murphy & Love
1950; Sampson et al. 1951). Introduced to the Hawaiian Islands about 1910 (Whitney et al. 1939; BISH
624327) and to New Zealand as early as 1840 (Zotov 1963), it is now well established in these regions. In
the United Kingdom, it has been reported as introduced with wool-waste (Lousley 1961; Ryves 1988), but
has not become fully naturalized (Stace 1997).

Darbyshire et al., The genus Rytidosperma in the U.S.A.

673

Plants densely to loosely caespitose, shortly rhizomatous. Ci
glabrous, usually scabrous immediately below the inflorescence, branching extravaginal, cataphylls scaly.
Leaves mostly basal, exceeded by or as long as the culms, flag leaf blades usually reaching the inflorescences;
sheaths glabrous or with scattered hairs, with apical tufts of hairs, hairs to 4 mm long; ligules 0.2-0.5
mm long; blades (5-)15-25(-75) cm long, to 2 mm wide, flat or involute, glabrous or pubescent. Inflores-
cences racemose or with a few branches, 5-15 cm long, lanceolate; rachis scabrous; pedicels shorter than
spikelets, scabrous. Spikelets (7-)10-13(-16) mm long, 6-7(-10) florets; glumes (7-)8-13(-16) mm long,
surpassing florets, subequal, lanceolate, subacute, glabrous or sometimes with a few hairs, light green to
stramineous often purple at margins and apex; lower glumes (5-)7-veined; upper glumes 5(-7)-veined;
rachilla segments 0.1-0.2 mm long; calluses (0.6-)0.9-1.5(-2) mm long, hairs 1.0-1.5 mm long and not
or barely reaching the lower lemma hairs; lemmas 2.5-3.5(-4.5) mm long, (7-)9 veins, lower row of hairs
dense, hairs not or just reaching the upper rows, upper row of hairs reaching or slightly exceeding the base
of the awn, scanty medial tufts usually present or sometimes absent, glabrous elsewhere; lobes 5-10 mm
long, abruptly aristate; awn 11-14 mm long, lightly twisted column 2-3 mm long, somewhat reflexed at the

or with a few hairs, veins ciliate; anthers 0.3-2 mm long. Caryopses 1.7-2.1(-2.5) mm x 0.8-1. 1(-1. 3)
mm; embryos 0.8-0.9 mm long; hila 0.4-0.5 mm long. 2n = 24; Australian plants (Abele 1959; Brock &
Brown 1961) and Californian plants (this paper).

This highly variable species is endemic to southern Australia where it “displays a bewildering variety
of forms” (Vickery 1956). Connor and Edgar (1979) described the upper hairs on the lemma as rarely form-

obtusatum (Benth.) Connor & Edgar, but all material from the United States is typical var. racemosum.

Naturalized primarily as a ruderal weed in disturbed areas less than 200 m in elevation. It is a common
weed of lawns and roadsides around Berkeley, California (Blumler 2001; Ertter, personal observation), and
has also been collected at Davis (Fig. 4D). St. John (1973) stated that it was introduced to Hawai'i in 1937,
but we were unable to locate specimens to confirm this.

Introduced and grown for forage trials at several locations in North America, this species is frequently
confused with R. penicillatum. Although the earliest North America record located dates from cultivation at
Berkeley in 1941, it was probably grown in earlier times under the name of Danthonia pilosa. It was present
in New Zealand as early as 1840 (Zotov 1963). In the United Kingdom, var. obtusatum has been reported
as introduced with wool-waste (Lousley 1961; Ryves 1988), but has not become fully naturalized (Stace
1997).

' South Wales Natl. Herb.

Vickery, J.W. 1 956. A revision of the Australian species of Danthonia DC. Contr. New
2:249-325.

WAGNEit W.L., D.R. Herbst, and S.H. Sohmer. 1 999. Manual of the flowering plants of Hawaii Revised edition. Bishop
Mus. Spec. Publ. 97: i-xviii + 1-1919.

Waters, C, B.G. Murray, G. Melville, D. Coates, A. Young, and J. Virgona. 201 0. Polyploidy and possible implications for
the evolutionary history of some Australian Danthonieae. Autral. J. Bot. 58:23-34.

Whitney, L.D., E.Y. Hosaka, and J.C. Ripperton. 1 939. Grasses of the Hawaiian ranges. Hawaii Agric. Exp. Sta. Bull.

Weintraub, F.C. 1953. Grasses Introduced into the United States. Agricultural Handbook No. 58. Forest Service,
U.S. Department of Agriculture, Washington, DC, U.S.A.

Zotov, V.D. 1963. Synopsis of the grass subfamily Arundinoideae in New Zealand. New Zealand J. Bot. 1:
78-136.

TYPIFICATIONS OF NAMES IN

AGALINIS, GERARDIA, AND TOMANTHERA (OROBANCHACEAE)

J.M. Canne-Hilliker

John F. Hays

8530 Wani Drive
Irvington, Alabama 36544, U.SA.
agalinisman@hotmail.com

When o

taxonomic treatment of Agalinis Raf. (nom. cons.) type specimens were not located for
md infraspecific taxa of Agalinis, Gerardia L. (sensu Pennell 1935) and Tomanthera Raf.
terial of A. Michaux, F. Pursh, T. Nuttall and C.S. Rafinesque was not found within her-

designated here for names for which no original material was located. The majority of neotypes were selected
from our own collections so that duplicates (isoneotypes) could be deposited in several herbaria. Care was
taken to ensure that new types are concordant with current usage of names as we understand them after
many collective years of scientific study of Agalinis. Each neotype conforms to the original morphological
description of the taxon and was collected from the geographical region indicated in the protologue. De-
scriptions of new species of Agalinis in Rafinesque’s New Flora of North America are particularly informative
and generally unambiguous.

In only one case was it not possible to identify the taxon to which a name applies although conflicting
attempts have been made by various authors. The name Gerardia erecta Walter ex J.F. Gmel, the basionym
of Agalinis erecta (Walter ex J.F. Gmel.) Pennell, is recommended for rejection under Art. 56.1 of the Inter-
national Code of Botanical Nomenclature (McNeill et al. 2006) as outlined in McNeill et al. (2007).

An illustration by Rafinesque was chosen as lectotype for a name published by him. Syntypes annotated
by Bentham, and initially selected as types by F.W. Pennell but not cited as lectotypes are designated here
in second step lectotypifications. Lectotypes were selected for a name published by A. Gray and a name
published by Rafinesque.

AgaUnis corymbosa Raf., New Fl. N. Amer. 2:63. 1837. Tm: U.S.A. Florida. Santa Rosa Co.: TIS. 28N. S.24, SE4SW4, ca.

100 plants in mesic to wet longleaf pine savanna, 10 Oct 1998, J.E Hays 2005 (neotype, designated here: NLU!; isoneotype: FLAS!).
Rafinesque’s ample description of plants he knew from “Carolina and Florida” describes well Agalinis pinetorum
var. delicatula Pennell. This a slender stemmed, narrow leaved variant of A. harperi Pennell.

Agalinis longifolia Raf., h

We agree with Pennell (1929) that Rafinesque’s morphological description of plan
streams New Jersey to Virginia” fits well the mid- Atlantic populations of A. purpure

Agalinis maritima var. gracilis Raf., New Fl. 1
purpurea, 30 Aug 1978,J.M. Canne and KA. Hrusch
Agalinis (originally misspelled Agalims) marit

Vmer. 2:62. 1837.

I (Raf.) Raf. var. g

I- BoL Res. Inst Texas 4(2): 677-681.

678

northern portion of the species range along the coast of the New England states.

i Raf., New FI. N. Amer. 2:62. 1837. l

Rafinesque described A.

'laritima var. pumila as “2 or 3 inches high.” Plants of this description are A. maritv
a the coastal regions of New England. Rafinesque knew the species only from “
England to Chesapeak bay.”

We agreed with Pennell (1920, 1929, 1935) who placed A. microphylla ]
Nutt. Rafinesque’s description of a Leconte collection from Florida in the
only to A. aphylla because he noted the deeply grooved and ribbed stem,
“spicate” inflorescence, short calyx lobes, and the few, alternate, virgate b
from Duval Co., Florida is from an area in which Leconte is known to ha

. in synonymy with A. aphylla
»llins herbarium is attributable
: very small, adpressed leaves,
ches. The collection by Curtiss
worked (Gray 1883).

Rafinesque’s mention of angular, filiform stems; sulcate, obtuse, callose (silicified), leaves; peduncles much
longer than leaves; minute, callose calyx teeth; and short broad corollas clearly denote the species as treated by
Pennell (1929, 1935). Rafinesque noted the species occurred in “West Tennessee, Alabama and Florida.”

Rafinesque published Agalinis palustris as a new name for Gerardia purpurea L. (= Agalinis purpurea (L.) Pennell)
(typ. cons.). Although A. palustris Raf. is an illegitimate name, in accordance with Art. 55.2 of the International
Code of Botanical Nomenclature (McNeill et al. 2006 ) the varietal names published by Rafinesque under
A. palustris are legitimate. Agalinis palustris var. corymbosa was described as “branches crowded corymbose.”
Rafinesque provided no information about the distribution of his varieties of A. palustris but noted that the
species occurred “From New England to Carolina.” He did not mention specimens seen in herbaria.

;s of A. palustri

Agalinis setacea var humUis Raf. New Fl. N Amer 2:64.

and Oken St. on NE side of Egg Harbor, sandy, weedy, | " -

Solidago.Uspedeza,7Sepl9
Rafinesque listed the range of Agalinis setacea as from “New Jersey t
described his varieties based on height; A. setacea var. humilis was s;

A. New JERSK Atlantic Co.: jet. Co. Rd. 563
Pinus woods with Heterotheca, Eupatorium.

3ama and Kentucky, Illinois” and

679

are of little taxonomic significance because both characters vary within populations. Also, as in other spe-
cies oiAgalinis, plants of A. setacea in more southern locations generally grow taller and have more branches
than those in the northern extremes of the species range.

Rafinesque described A. temifolia var. humilis as “semipetal few branches,” a minor distinction of no taxonomic
value for plants of this very widespread and variable species. Although Rafinesque listed the range of the
species “from Canada to Florida and Missouri” he did not publish locality data for his new varieties.

Pennell (1929) was unable to locate a type for Agalinis virgata Raf., but nonetheless used the name for a taxon
we consider to be a taxonomically insignificant variant of A. purpurea (L.) Pennell. Rafinesque described
plants from “glades of Pine woods in South New Jersey near Mullica Hill.”

lanthera Raf. Florula ludov. 50. 1817. '

Caddo Parish: along LA 789 S of Spring Ridge and

Rafinesque’s description clearly refers to a variant of Agalinis tenuifolia (Raf.) Raf. in which the style is held
above the densely white lanose abaxial anthers. The plants were described as three feet tall, but plant height
is variable within and among populations. The description by Robin (1807) upon which Rafinesque based
his account of G. kucanthera is an excellent depiction of A. tenuifolia.

i Raf. Med. Repos. II. 5:361. 1808. '

We and Pennell (1929, 1935) were unable to locate original Rafinesque specimens referable to G. maritima
in North American and European herbaria. However, the plate of Gerardia maritima by Rafinesque at NY is
an identifiable portrayal of the taxon. Pennell (1929) stated that “An unpublished plate of Rafinesque’s in the
library of the New York Botanical Garden denotes clearly the plant here considered.” Merrill (1949) noted
that the plates at NY are likely the only remaining evidence that illustrate Rafinesque’s American species
prior to the loss of Rafinesque’s herbarium in a shipwreck off Long Island in 1815. Gerard (1885) quoted the
notation made by Rafinesque on the first plate of the set in which Rafinesque explained that the plates were
proofs of plates lost in the shipwreck of 1815, and are figures of plants for which names were published in
1807, 1808 and 1814. Rafinesque’s plate of Gerardia maritima shows a complete plant and nine numbered
insets which include in numerical order: a leaf; flower; calyx and style; calyx opened showing the ovary with
style; longitudinal section of a corolla; capsule with calyx; lateral view of an open capsule; cross section of a
capsule; and seeds. The habit sketch shows an elongate main stem with shorter basal branches, and flowers
with blunt calyx lobes that are distinctive of Agalinis maritima (Raf.) Raf. in the northern part of its range.
Tomanthera lanceolata Raf. , New Fl. N. Amer. 2:66. 1837 . Type: Erinus, Gerardia aunculata, Muhlenbcig Herbanum 851

Rafinesque (1837) cited “ Erinus! africanus Muhl.” with his description of T. lanceolata

1. Earlier, Nuttall (1818)

ndGerardia

681

sonville, Louisiana” (first step typification). However, in his protologue Bentham listed two collections,
“Jacksonville and Louisiana, Drummond,” for his new taxon. There is a specimen at Kew attributed to Drum-
mond from Louisiana upon which Bentham wrote “G. temifolia leptophylla." This specimen, the lectotype, is
a plant of Agalinis gattingeri (Small) Small and is annotated with this name by Pennell.

a Benth., Companion Bot. Mag. 1:209. 1

Bentham cited “Jacksonville and St. Louis, Drummond” for specimens of G. temifolia var. macrophylla.
Pennell (1920) reported that he had seen a “Fragment of type, from Kew Herbarium, labeled St. Louis.”
From the two syntypes he thus selected the St. Louis specimen, via the fragment, as the lectotype (first step
lectotypification).

ACKNOWLEDGMENTS

We thank J. Dorfman, The LuEsther T. Mertz Library (NY), for the digital image of the plate by Rafinesque;
A. Freire-Fierro (PH) for the digital image of the type of Tomanthera lanceolata, C. Nepi (FI), and V. Fonjal-
laz and L. Gautier (G-DC) for sending digital images of specimens at their institutions; L. Amadei (PI), S.E.
Began (DWC), S. Marner (OXF), G. Reid (LIV), O. Ryding (C), A. Smith (E), L. Wolstenhole (MANCH) for
their efforts in searching for specimens; K. Gandi (GH) for answers to questions regarding nomenclature.
We also thank T. Lammers for a very helpful review of our manuscript. Canne-Hilliker thanks the curators
at ACAD, CAN, DAO, FSU, GA, GH, K, MIN, NCU, PH, TRT, USCH, and WIS for loan of specimens. Hays
thanks the curators at AKG, AUA, BRIT, DUKE, ELAS, FSU, GA, IBE, LAF, LSU, LTU, MISS, MO, NLU, NO,
ODU, SWSL, UNA, UAM, UMO, NCU, USAM, US, USF, TENN, UWEP, and VDB for loan of specimens and

REFERENCES

Bird, CD. 1967. The mosses collected by Thomas Drummond in Western Canada, 1825-1827. Bryologist
70:262-266.

Gray, A. 1883. Some North American botanists. IV. John Eatton LeConte. Bot. Gaz. (Crawfordsville) 8:197-199.
Gerard, W.R. 1885. Reliquiae Rafinesquianae. Bull.Torrey Bot. Club 12:37-38.

McNeill J., F.R. Barrie, H.M. Burdet, V. Demoulin, D.L Hawksworth, K. Marhold, D.H. Ncoison, J. Prado, RC. Silva, j.E. Skog, J.H.
WiERSEMA, AND NJ.Turland (eds.). 2006. International code of botanical nomenclature (Vienna Code) adopted by
the Seventeenth International Botanical Congress Vienna, Austria, July 2005. Regnum Veg. 146.

McNeill, J„ S.A. Redhead, and J.H. Wiersima. 2007. Guidelines for proposals to conserve or reject names. Taxon
56:249-252.

Merrill E.D. 1 949. Index Rafinesquianus. Arnold Arboretum, Jamaica Plain, Massachusetts.

Merrill, E.D. and S-Y. Hu. 1 949. Work and publications of Henry Muhlenberg, with special attention to unrecorded
or incorrectly recorded binomials. Bartonia 25:1-66.

NuttallT. 1818. The genera of North American plants, 2. Facsimile of the 1818 edition, Hafner Publ. Company,
New York. 1971.

Pennell, F.W. 1920. Scrophulariaceae of the southeastern United States. Proc. Acad. Nat. Sci. Philadelphia
71:224-291.

Pennell RW. 1 929. Agalinis and allies in North America. II. Proc. Acad. Nat. Sci. Philadelphia 81:111 -249.

Pennell, F.W. 1935. The Scrophulariaceae of eastern temperate North America. Monogr Proc. Acad. Nat. Sci.
Philadelphia 1:1-650.

Robin, C.C. 1807. Voyages dans I'interieur de la Louisiane

? 3:313-551. Paris.

682

BOOK REVIEW

A NEW COMBINATION IN LOLIUM PERENNE (POACEAE: POEAE);
L. PERENNE SUBSP. STOLONIFERUM
Joseph K.Wipff,lll

West Coast Research Center
Barenbrug USA, Inc.

36030 Tennessee Rd.

Albany, Oregon 97322, U.S.A.

ABSTRACT

RESUMEN

Lawson (1836) described a very distinct taxon of the Lolium perenne complex as Lolium perenne var. stoloniferum,
giving it the common name of ‘Spreading Ryegrass.’ Lawson described this new taxon as follows;

n very sparingly.

The very long determinate-stolons, which root at the nodes, and aggressive spreading habit of L. stoloniferum
are very distinct from L. perenne sensu stricto and warrants recognition at the subspecific level, thus neces-
sitating the following new combination.

le L. subsp. stoloniferum (C. Lawson) Wipff, comb, et stat. i
C. Uwson, Agric. Man. 104. 1836. Type: OREGON. Linn Co.: S of Hwy 226 anc
Lebanon, on the West Coast Research Center farm, Barenbrug USA, Inc. 22 Jun 2i
here; isoneotypes: BRCH, BRIT, L, MO, NY).

specimens cited within the protologue and no record of collecti

Discussion regarding the basionym:

At first it appears that Lawson (1836) was describing a new species because his protologue (pg. 104) started
as follows; “10. Spreading Ryegrass (L. stoloniferum).”

But an examination of his treatment reveals several places where his intention that he recognized this
taxon at the varietal rank, and not at a specific rank. Indicating he was actually describing a new variety of
L. perenne and not a new species of Lolium. Lawson (1836) explicitly recognized this new taxon at the rank
of a variety on the following pages.

1. Pg. 102. . . .“Uke other plants which have received an extensive cultivation, there are several varieties of the

Common Ryegrass, the principal of which are as follows;— ...” ^

He then goes on to describe 11 varieties of L. perenne, one which (no. 10) being Spre^ing E^gi^s.

2. Pg. 104. “10 Spreading Ryegrass (L stolom/erum).-This name is applied to a remarkably stoloniferous

variety, the seeds of which were procured from Germany, ...”

3. Pg. 105. “The above are the most esteemed varieties of Lolium perenne, but there are many more of infenor

importance, and possessing less permanent characters.”

684

ACKNOWLEDGMENTS
i (HUH) for his invaluable as«

REFERENCES

Lawson, C 1836. Agriculturist's manual: agricultural plants cu
a report of Lawson's Agricultural Museum in Edinburgh. \

tivated in Europe; climate of Great Britain. Forming
lliam Blackwood and Sons, Edinburgh, Scotland.

PALEOCHARIS Nl

:N. and SR NOV (CYPERACEAE) IN
CANADIAN AMBER

George O. Poinar, Jr.

The fossil is represented by a mature achene wit!
perianth bristles. The specimen is complete, wi
Sion and distortion occurred during the fossili:

Paleocharis Poinar & D.J. Rosen, gen. nov. Type: Paleocham nearctka Poinar & D.J. I

DESCRIPTION

/ith an attached stylopodium (tubercle) and numerous hair-like
e of the bristles detached. Some minor compres-

Generic diagnosis: Achenes with persistent style base (stylopodium), and numerous (over 200) smooth
hair-like bristles equaling the length of the achene plus the stylopodium. The proximal half of each bristle
is composed of thick-walled dark cells while the distal half is formed by thin-walled, light-colored cells.

Poinarand Rosen, F

687

Paleocharis nearctica Poinar & DJ. Rosen, sp. nov. (Figs. 1-4). '

Specie diagnosis: Flowers with numerous (over 200) smooth, hair-like perianth bristles, 2.6-3.0 mm long,
equaling the length of the achene plus the stylopodium (Figs. 1, 2); each bristle composed of a single row of
14-20 elongate cells; cells 0.1-0.26 mm long x 0.1-0.16 mm wide; proximal half of each bristle composed
of thick-walled dark cells while distal half formed by thin-walled, light-colored cells (Fig. 3); stamens not
seen; style with short lobes. Achenes narrowly oblong, lenticular, 1.7 mm long X 0.48 mm wide; surface with
transversely oblong cells, smooth except for some delicate microscopic scales ranging from 0.58-0.7 mm

lanceolate-obclavate, 1.2 mm long X 0.28 mm wide, dark brown (Fig. 2).

Etymology. — “Paleo” is from the Greek “palaios” for ancient and “charts” is Greek for beautiful, “nearctica”
refers to the geographic location of the fossil.

DISCUSSION

Consideration was given to the possibility that the achene could belong to other plant taxa. Members of
the Asteraceae also have achenes as fruits, and a persistent calyx (pappus) comprising numerous capillary
bristles in many species (Zomlefer 1994; Simpson 2006). However, the bristles are positioned apically usu-
ally crowning the achene rather than basally as in Cyperaceae (Zomleferl994; Simpson 2006). Members
of Salix L. and Populus L. of the Salicaceae have seeds with long, silky hairs, however these seeds do not
contain tubercles or have pointed tips and the hairs on the seeds are flexible and single-celled, not stiff and
multicelled. In addition some of the hairs on Salix and Populus seeds arise from the seed coat, whereas in
the fossil, all of the bristles are hypogenous (Woody-Plant seed Manual 1948).

Paleocharis shows morphological resemblance to extant species in Eleocharis R. Br. and Rhynchospora

numerous elongate, smooth perianth bristles (Flora of North America Editorial Committee 2002). However,
the narrow, elongate achene and stylopodium and numerous smooth, stiff, bi-colored bristles comprised of
a single row of cells in Paleocharis are unique characters not known to occur in extant sedges. A comparison
of the qualitative characters of the fossil fruit with those of the extant genera mentioned above is shown in
Table 1. As Paleocharis overlaps in more achene morphological characters and dimensions with Eleocharis and
Rhynchospora, we suggest its placement as an extinct member of Cyperoideae Suess (Simpson et al. 2007).

The darker, heavier cells forming the basal portion of the bristles may have insured that the achenes
landed with the base making initial contact with the substrate and provided for the absorption of heat to
bring about germination as suggested by Lye (2000) for some temperate species of Cyperaceae. The pres-
ence of numerous smooth perianth bristles suggests that P. nearctica achenes were wind-dispersed, similar
to those of Eriophorum. In contrast, Eleocharis and Rhynchospora, both of which a predominance of species
with variously barbed bristles, are known to be dispersed by animals, especially birds (Sauer 1988). While
most dispersal is external on feathers, there are records of some Eleocharis seeds carried in the digestive
system of birds (Sauer 1988).

Fossil sedge fruits are fairly common in Tertiary deposits (Collinson et al. 1993; Smith et al. 2009) but
there are no accepted Mesozoic fossils. Putative Cyperaceae pollen was reported from Cretaceous deposits in
New Zealand (Couper 1953), Berry (191 1) described Carex clarkii as a putative leaf fossil from Late Cretaceous
Coniacian deposits in New Jersey and Caricopsis laxa Samylina was described as a putative leaf fossil from
the Early Cretaceous of Siberia (Samylina 1960). While Collinson et al. (1993) cite C. laxa as the earliest
sedge fossil, they comment, “We know of no well-substantiated leaf fossils of the family” and others have
regarded C. laxa as not being Cyperaceae (Daghlian 1981; Friis et al. 1987). Pollen from the Lattest Santonian
to Maastrichtian strata in Western Canada described as Penetetrapites inconspicuus Sweet, was later considered
as possibly belonging to the Cyperaceae (Sweet 1986; Braman & Koppelhuis 2005) but this has not been
confirmed. More recently, Goetghebeur (1998) and Smith et al. (2009) reported that fossils of Cyperaceae
are only known with certainty from the Paleocene.

Journal of the Botanical Research Institute of Texas 4(2)

The climate in Alberta during the Campanian Stage of the Late Cretaceous was subtropical to warm
temperate. A large epicontinental sea divided North America and the resin-producing araucarians were not
far from the Western bank of that sea (Braman & Koppelhuis 2005; Poinar & Poinar 2008; Smith et al. 1994).
Based on chemical analysis, Canadian amber from Alberta was produced by araucarian trees, presumably
belonging to the genus Agathis, commonly known as Kauri (Lambert et al. 1990). In the Waipoua araucarian
forest of New Zealand, which is the only remaining virgin Kauri forest in the world, sedges, especially Gahnia
xanocarpa (Hook.), are one of the dominant features of the undergrowth (Cockayne 1908; McGregor 1948). It is
possible that Pakocharis nearctica was a common sedge in the undergrowth of the Canadian araucarian forest.

Journal of the Botanical Research Institute of Texas 4(2)

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Braman, D.R. and E.B. Koppelhuis. 2005. Campanian Palynomorphs. In: PJ. Currie and E.B. Koppeihus, eds. Dinosaur Pro-
vincial Park, A spectacular ancient ecosystem revealed. Indiana University Press, Indianapolis. Pp. 101-130.

BREMEf^ K. 2002. Gondwanan evolution of the grass alliance of families (Poales). Evolution 56:1 374-1 387.

Cockayne, L. 1 908. Report on a botanical survey of the Waipoua Kauri Forest. John Mackay, Govt. Printer, Wellington.

COLLINSON, M.E., M.C. Boulter, and P.L Holmb. 1 993. Magnoliophyta ('Angiospermae'). In: MJ. Benton, ed.The Fossil
Record-2. Chapman & Flail, London. Pp. 809-841 .

CouPER R.A. 1953. Upper Mesozoic and Cainozoic spores and pollen grains from New Zealand. New Zealand
Geol. Survey Palaeont. Bull. 22:1-75.

Daghlian, R.M.T. 1981. A review of the fossil record of Monocotyledons. Bot. Rev. 47:5 17-555.

Eberth, D.A. and A.P. FIamblin. 1 993. Tectonic, stratigraphic and sedimentological significance of a regional dis-
continuity in the Judith River Group (Belly River Wedge) of southern Alberta, Saskatchewan and northern
Montana. Canad. J. Earth Sci. 30:1 74-200.

Flora of North America Editorial Committee. 2002. Flora of North America north of Mexico, Vol. 23. Magnoliophyta:
Commelinidae (in part): Cyperaceae. Oxford University Press, New York.

Friis, E.W., W.G. Chaloner, and P.R. Crane (eds.). 1 987.The origins of angiosperms and their biological consequences.
Cambridge University Press, Great Britain

GoETGHEBEua P. 1 998. Cyperaceae. In: K. Kubitzki, H. Huber, PJ. Rudall, PS. Stevens, andT. Stutzel, eds. The families
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COLEATAENIA GRISEB. (1879): THE CORRECT NAME FOR
SORENGIA ZULOAGA & MORRONE (2010) (POACEAE: PANICEAE)
Robert J. Soreng

National Museum of Natural History
Smithsonian Institution
Washington, DC 20013-7012, U.S.A.

The new genus Sorengia Zuloaga & Morrone, proposed in Zuloaga et al. (2010), is illegii
McNeill et al. 2006), in that the type species of the genus Coleataenia Griseb. (1879) was in
of the species of the new genus as a taxonomic synoriym. Coleataenia gynerioides Griseb. (
as a synonym of Sorengia prionitis (Nees) Zuloaga & Morrone. Twelve new combinations
accepted in Sorengia by Zuloaga et al. (2010) are proposed here.

. Konigl. Ges. Wiss. Gottingen 24(1);308

Coleataenia caricoides (Nees ex Trin.) Soreng, c

Coleataenia longifolia (Torr.) Soreng, comb, no^

Coleataenia longifolia subsp. abscissa (Swallen) Soreng, comb. nov. Basionym: i

Coleataenia longifoUa subsp. combsu (Scribner & C.R. Ball) Soreng, comb, r
& C.R. Ball, Bull. Div. Agrostol., U.S.D.A. 24:42. 1901.

Coleataenia longifolia subsp. elongata (Scribn.) Soreng, comb. nov. BAsiot
Coleataenia longifoUa subsp. rigidula (Bose ex Nees) Soreng, comb. nov. Ba

Coleataenia petersonii (Hitchc. & Ekman) Soreng, comb, i

Coleataenia prionitis (Nees) Soreng, comb. nov. Basionym; ]

Colea

REFERENCES

McNeill, J„ F.R. Barrie, H.M. Burdet, V. Demoulin, D.L. Hawksworth, K. Marhold, D.H. Nicolson, J. Prado, P.C. Silva, J.E.
Skog, J.H. Wiersema, and NJ.Turland (eds). 2006. International code of botanical nomenclature (Vienna Code).
Regnum Veg.146. A.R.G. Gantner Verlag KG.

Zuloaga, F.O., M.A. Scataglini, and O. Morrone. 2010. A phylogenetic evaluation of Panicum sects. Agrostoidea,
Megista, Prionita and Tenera (Panicoideae, Poaceae): Two new genera, Stephostachys and Sorengia. Taxon
59:1535-1546.

CONGRUENCE BETWEEN ALLOMETRIC COEFFICIENTS AND PHYTOGENY
IN STIPOID GRASSES: AN EVO-DEVO STUDY
Jack Maze

RESUMEN

INTRODUCTION

The idea of a relationship between evolution and development is as old as evolution itself (Gilbert 2003) even
used by Darwin (1859) as evidence for unity of type (Gilbert 2003). In the middle of the 20th century the
rise of the Modern Synthesis resulted in genetics supplanting development as an explanatory phenomenon
and it was argued that genetics and not development held the key to evolution (Gilbert 2003). In spite of
genetic ideas coming to dominate in evolutionary studies, a relationship between development and evolu-
tion was not abandoned. For example, evolution, as descent with modification, formed the conceptual basis
in attempts to understand the basic structure of the flower (e.g., Barnard 1957, 1960; Tepfer 1953; Tucker
1959) or of vascular plants in general (Meeuse 1966). Gould’s Ontogeny and Phytogeny (1977) emphasized the
relationship between the two phenomena and stimulated studies wherein evolutionary change was described
in terms of modified allometries (see e.g., Gibson & Diggle 1997). There were also attempts to link evolution
and development through emphasizing the developmental changes that have occurred with evolution (see
McMahon & Hufford 2002; Olson 2003; Richards et al. 2006 and references therein). Still another approach
to linking development and evolution described the changes in relative frequency of histogenetic events, e.g.,
cell division and enlargement, that occur with evolution (Kam &r Maze 1974; Maze et al. 1972; Stebbins 1967).

J.BotR«s. Inst Texas 4(2): 693 -7

Journal of the Botanical Research Institute of Texas 4(2)

Three developments in biology led to a renewed interest in the relationship between evolution and
development (Gilbert 2003). One was the ability to infer more precise phylogenetic relationships based on
numerical analyses of molecular data (Soltis et al. 2000). The second was the identification of genes involved
in the development of organisms. Once these genes were known mutations in them could be used to deter-
mine their role in developmental processes, such as the nature (see Friedman et al. 2004; Meyerowitz 2002)
or positioning (Smith et al. 2006) of appendages. In addition information was obtained on how genetically
mediated changes affected growth rates which, in turn, are expressed phenotypically as changes in size and
shape (Coen et al. 2004; Langlade et al. 2005; Rolland-Lagan et al. 2005). The third development resulted
from re-evaluation of basic precepts and led to a conclusion of “..the inability of the neoDarwinian synthesis
to account for many phenomena of higher-level phenotypic organization” (Muller & Newman 2005a). Similar
arguments were presented in Maze & Finnegan (2008).

Modern studies in evolutionary development (evo-devo) include those which address the incorpora-
tion of developmental traits, either structural (see Olson 2003; Friedman et al. 2004) or molecular (Arendt
2003) in established phylogenetic trees or the genetic changes underlying adaptation (Hoekstra & Coyne
2007). Evo-devo arguments have also been used to explain the origin of novelty as being the result of
environmentally induced developmental events (Muller & Newman 2005b; West-Eberhand 2005) that
become incorporated into the DNA. Jablonka & Lamb (1995) and Steele et al. (1998) have also argued for
the incorporation of environmentally induced traits. Pigliucci (2007), as well, has raised the question of the
necessity of an extended evolutionary synthesis which incorporates environmental changes more directly
into accounts of evolutionary change.

Another series of studies linking evolution and development are those seeking a common underlying
cause in this case in non-equilibrium thermodynamics and information theory (Maze 1999; Maze et al. 1990,
2001a, b, 2002, 2003a,b’ 2005; Robson et al. 1993). In these studies it was reasoned that the morphological
changes that occur with both evolution and development are the result the transformation of matter, i.e.,
the production of information, through which energy dynamics are carried out. This is seen, for example,
in the production of high energy compounds such as ATP and NADH or carbohydrates, cellulose, secondary
metabolites and proteins. The basic argument is that as matter is transformed during development it can be
understood as information, “in-formed matter”. This new morphological organization of information becomes
part of the totality of information that delineates a species, i.e., its information system (Brooks, 2001, 2002;
Brooks & Wiley 1988; Maze et al. 2005). This results in the expansion of the information system due to
matter transformation and that, along with the addition of new information through genetic mutation and
recombination, results in an increase in the complexity of the information system of a species. Once the
information system of a species reaches a certain stage of complexity it bifurcates, expressed biologically
as speciation (Brooks. 2001; Brooks & Wiley 1988). This view of speciation, admittedly a unique one. has

Here 1 take a different approach to the study of evolution and development, a comparison of the rela-
tionships inferred from a phylogeny with the relationships inferred from an analysis of growth phenomena
represented by allometric coefficients. There is no doubt that there is some sort of relationship between
allometric coefficients and evolutionary change (Coen et al. 2004; Langlade et al. 2i

2005); see also Gould (1977). But, what is the nature of that relationship beyond tl

in their allometric coefficients? Specifically 1 explore the idea there is some sort of predictability between
allometric coefficients and phylogeny. In other words, by knowing one, e. g., allometric coefficients, can
some sort of predictive statement be made about phylogeny? A predictive relationship between allometric
coefficients and phylogeny is of interest as it may indicate some deep-seated underlying cause such as was
argued by Maze (1999); Maze et al. (1990, 2001a,b, 2002, 2003a,b> 2005) and Robson et al. (1993).

MATERIALS AND METHODS

Plants. The plants used in this study are grasses in the genus Achnatherum, tribe Stipeae (Poaceae; Pooideae).

)5; Rolland-Lagan et a

lers by acronym for each collection, number of individuals measured.

LEMENT1 -8. 28.3 km n Enterprise on Oregon State H ighway 3, Wallowa Co., OR, U. S. A.; plants growing in forest of ponderosa
pine and Douglas fir. 45.80 N, 11 7.21 W. 9 Jul 1998

LEMC0L01 -8. just across Kittitas-Chelan Co. line in Kittitas Co. along Colockum Pass Road, WA, U. S. A.; plants growing in an
open stand of ponderosa pine. 47.50N, 1 20.1 9W. 1 3 Jun 2002.

LEMFOX1 -8. 7 km from U. S. Forest Service Road 4240 on road 200, Crook Co., OR, U. S. A.; plants in deep soil in among pon-
derosa pine. 44.1 6N, 120.10W. 15 Jun 2002.

LEMIND1-8. 15.8 km s. Foothill Road on Indian Springs Road, Twin Falls., Co., ID, U. S. A.; plants growing among shrubs in
rolling hills. 42.34N, 1 14.57W. 1 1 Jun 2004.

LEMSIE1-8. 2.4 km. s. of Graeagle on Calif St. Highway 89, Plumas Co., CA, U. S. A.; plants growing with ponderosa pine.
39.45N,120.37W.18Jun2004.

LEMCLR1 -6. 1 km. W. of Calif St. Highway 89, On Clark Creek Road, Shasta Co., CA, U. S. A.; plants growing with Quercus kellogii
and ponderosa pine. 41 .1 5N, 1 21 .72W. 1 9 Jun 2004.

Achnatherum hendersonii

HENCOL1 -8. along Tarpescan Creek Road where it joins Colockum Pass Road, Kittitas Co., WA, U. S. A.; plants growing in shal-
low soil in sparse vegetation. 47.47N, 1 20.20W. 1 3 Jun 2002.

HENFOX1-8. 7 km from U. S. Forest Service Road 4240 on road 200, Crook Co., OR, U. S. A.; plants growing in shallow soil in
sparse vegetation. 44.30N, 120.20W. 15 Jun 2002

WALBON1-8. near Boner Springs, middle sw ’A of se ’4 of section 24,T3N, R45E., Wallowa-Whitman Nat. For., along FS Road
46, Wallowa Co, OR, U. S. A.; plants growing in shallow soil in sparse vegetation. 45.67N X 1 1 7.1 3W. 26 Jun 1 993.
WALSK01-8. middle of section 9, T1 2S, RISE, Ochoco N. F, Crook Co., OR, U. S. A.; plants growing in shallow soil in sparse
1. Site 3001 1 USDA Forest Service R-6 Threatened, Endangered and Sensitive Plant Surveys. 44.53N X 1 20.60W,

28, 29 Jun 2003.

ELMSIE1-10. 2.4 km. s. of Graeagle on Calif St. Highway 89, F
39.45N, 120.37W. 1 Jun 2001.

ELMBUL1-8. growing along old road just n. of Bull Mountain near g

Forest Service Road 27, Ochoco National Forest, Crook Co. OR, U. S. A. 44.5 1 N, 1 20.60W. 1 5 Jun 2002.

ELMCOL1-8. along west bank Columbia River just north of bridge carrying U. S. Highway 395 across the Columbia River,
Stevens Co., WA, U. S. A.; plants growing with ponderosa pine. 48.63N, 1 18.13W. 18 Jun 2002.

ELMMON1-8. n. side Power House Road, just across U. S. Highway 395 on the west bound extension of Calif St. Hiway 167,
Mono Co., CA, U. S. A.; plants growing in sage brush. 38.05N, 1 1 9.1 7W. 1 7 Jun 2004.

ELMOLD1-8. 1 .6 km sw junction California State Highways 44 and 89 on 89, near Old Station, Shasta Co., CA, U. S. A. 40.68N,
121.30W.19Jun 2004.

IS Co., CA, U. S. A.; plants g

e guard on Forest Service Road 2730, 2 r

NELWL1 -1 4. from 1 . 1 km w. of Highway 97 on White Lake Road, s. of Pentiction B. C, Canada; plants growing open area with
ponderosa pine. 49.42N, 1 1 9.64W. 8 Jun 2001 .

NELCOL1 -8. from along west bank Columbia River just north of bridge carrying U. S. Highway 395 across the Columbia River,
Stevens Co., WA, U. S. A.; plants growing with ponderosa pine. 48.63N, 1 1 8. 1 3W. 1 8 Jun 2002.

NELENT1-3. from 28.3 km n Enterprise on Oregon State Highway 3, Wallowa Co., OR, U. S. A.; plants growing in forest of
ponderosa pine and Douglas fir. 45.80 N, 1 1 7.21 W. 9 Jul 1 998

NELANA1 -8 A nelsonii from rest stop on Anacharist Mt. Along Highway 3, British Columbia, Canada. 49.02N, 1 1 9.37W. Jun 2002.
NELMAN 1 -8 growing in gravel patch in parking lot a
B. C., Canada. 49.06N, 1 20.75W. 3 Jul 2004

since the direct assessment of time demands destructive sampling. Second, this would be extremelhy time
consuming since the preparation of spikelets of different ages requires the production of a large number of
microscope sections. Third, the lack of synchrony between developmental stages in the Stipeae (Maze et al.
1971, 1972) would introduce a problem in comparing spikelets of different ages. Fourth, the inability of get
sections that could be easily measured means that far less data could be gathered.

Maze, Allometric coefficients and phytogeny in stipoid grasses 697

In order to make comparisons between development, as represented by allometric coefficients, and
phylogeny, as represented by a phylogenetic classification I had to represent each species using allometric
coefficients as variables. That is easily done, do the required PCA for each species, take the first eigenvector
and then transpose it from a vector into a row of five variables where each variable, say first glume length, is
represented by its allometric coefficient. But it would be inappropriate to do a single PCA for each species.
Such would result in a data set with only five cases, one for each species, and five variables, the allometric
coefficients for the five features measured. Such a small data set is of little use in a comparison with the
results of a phylogenetic analysis since it is too small of a data set to produce meaningful results. Thus, to
generate a population of allometric coefficients for each species, I randomized all the spikelets within each
species, divided that randomized data set into groups of 94 spikelets each and did a PCA on each of those
groups. This meant that there were from 8 PCAs, for A. hendersonii and A. wallowaense, 20 for A. occidentale,
21 for A. nelsonii and 23 sets for A. lemmonii. Each group of 94 was checked to assure that all individuals
and populations collected for each species were included within it. A group size of 94 was chosen to assure
analytical stability, i.e., that the results were not an artifact of small sample size.

The randomization of spikelets was a choice made on developmental considerations. Allometric coef-
ficients for any set of spikelets are a numrical summary of the developmental events of the spikelets in that
set. Those developmental events will be the result of the genomic instructions for development and the
interactions between that genome and the environment it experiences during development. That environ-
ment has both external and internal components. The external environmental factors producing an effect
would be the likes of the continually changing day length, temperature, moisture, soil and neighboring
organisms that a developing plant experiences. The internal environment is established by the distribution
of growth promoting and inhibiting substances, e.g., hormones. The complexity of the internal environment
can be traced to the continually changing sources of growth effecting substances as growth centers appear
and disappear. The purpose of data randomization was to neutralize the effect of the genotype of any one
individual as well as any environmental effect on calculated allometric coefficients. Each 94 spikelet sample
included spikelets from all individuals and populations. Thus, any one of the sets would not have spikelets
that have all been subjected to similar environmental, both internal and external, or genetic effects. This
lowers the probability that any one PCA was biased because of an asymmetric distribution of environmental
influences.

As a result of the randomization within each species, subdividing of the data for each species into
groups of 94 spikelets and submitting each group to PCA, 1 generated an 80 x 5 matrix. Each of the 80 cases
represents the results of one PCA of the randomized 94 spikelets and the five variables were the allometric
coefficients i.e., the elements in first eigenvector for that PCA. The allometric coefficients were compared
with each other using the Kolomogorov-Smirnov test of variables. This is a test to determine if two variables
have a similar distribution as based on a distance function. This distance statistic was used to evaluate the
relationship among the allometric coefficients, a small distance indicating a similar distribution is taken as
evidence the allometric coefficients for the variables are similar. The similarity and differences among the
allometric coefficients were then evaluated by relying on the ontogenetic events whereby the different vari-
ables develop. Variables with similar allometric coefficients would be expected to show similar ontogenetic
events, e.g., patterns of cell division and maturation in comparable tissues.

Phylogenetic analyses. The phylogenetic analyses were based on 19 variables of both vegetative and
reproductive features (Table 2). To avoid analytical redundancy variables that described the features similar
to those used to calculate allometric coefficients were not included in the phylogenetic analysis. Although
there were only five species subjected to phylogenetic analysis, I wanted to be sure the phylogenetic signal
was strong, i.e., that the data describing the five species was sufficiently stable to give the same results re-
gardless of outgroup. To that end, four species were used as outgroups, Hesperostipa comata (Trin. & Rupr.)
Barkworth, Nasella viridula (Trin.) Barkworth, Achnatherum lettermam (Vasey) Barkworth and A. hymenoides
(Roem. & J.A. Schult.) Barkworth. The first two species are, like Achnatherum, in the Stipeae and, at one

699

by the species in the matrix of allometric coefficients. The dummy variable that recognized the lineage

species in were positioned in the data matrix of allometric coefficients. Where the remainder of the species
were positioned in the data matrix of allometric coefficients that dummy variable was given a value of 1.
The value for the dummy variable used to represent the lineage consisting of A. occidentak and A. nehonii
was established in the same way; that variable was coded as 2 where those species were positioned in the
data matrix of allometric coefficients and as 1 for the position occupied by the remainder of the species, A.
lemmonii, A. hendersonii and A. wallowaense. The third dummy variable that represented the lineage consist-
ing of A. lemmonii, A. hendersonii, and A. wallowaense was coded in the same way; it was given a value of
two where those three species were positioned in the data matrix of allometric coefficients and a value of 1
elsewhere. An example of this matrix of dummy variables along with the species names is in Table 3.

Comparison. The first step in comparing the allometric coefficients with the phylogenetic classifica-
tion was to summarize the data with principal components analysis (PCA). One PCA was done on the 80
X 5 data matrix of allometric coefficients and the other on the 80 x 3 matrix of dummy variables used to
describe the phylogenetic classification. The results of the two PCAs were compared with a Spearman rank
correlation coefficient. Only first PCA axes were compared since they are the best descriptors of the data. This
approach gives a single number, statistical in nature, summarizing the relationship between development
and evolution. A Spearman rank correlation coefficient was chosen because all the numerical manipulations
I used made me leery of using a parametric statistic.

All analyses were done using SYSTAT 4.0 (Wilkinson 1991).

RESULTS

Table 4 presents a comparison of the allometric coefficients for the five species as a Kolomogorov-Smirnov
test of variables. This statistic is a distance measure evaluating the distributions of the variables being tested.
The distributions for the allometric coefficients for glume length have a distance that is not statistically
significant. The distributions for all other pairs of variables have distances that are statistically significant.

linked to their developmental history. I also did a Spearman rank correlation on the allometric coefficients.
That is not shown but gave comparable results, the allometric coefficients for the lengths of the two glumes

The Spearman rank correlation coefficient between the PCA axis scores summarizing the allometric
coefficients and the PCA axis scores summarizing the phylogenetic classification is .790, p««.001; the
allometric coefficients and phylogenetic classification are strongly congruent, they are giving similar signals.
As a test of this approach I produced four other matrices of dummy variables describing classifications dif-
ferent from that inferred by the phylogenetic classification. The Spearman rank correlation between those
four alternate classifications and the allometric coefficients were 0.030, -0.137, 0.579 and 0.413, all lower
than the original test.

DISCUSSION

The similarity between the allometric coefficients for the length of the two glumes is hardly surprising.
Although not all the species included here have been studied developmentally, those that have, A. hender-
sonii (Mehlenbacher 1970) and A. lemmonii (Maze et al. 1972), show very similar patterns in initiation and
growth of the glumes. As well, the glumes of other Stipeae in which development has been described (Maze
et al. 1971; Kam 1974; Kam & Maze 1974) are like A. hendersonii and A. lemmonii. The glumes are little more
than acute to acuminate flaps of tissue without striking cellular differentiation in them other than relatively
simple epidermis, parenchyma and vascular tissues.

The dissimilarity in allometric coefficients for all the other spikelet structures measured is, likewise,
not surprising. Floret length, as measured here, is developmentally complex. One aspect of floret length.

Achnatherum lemmonii (23)
Achnatherum hendersonii (8)
Achnatherum wallowaense (8)
Achnatherum ocddentale (20)
Achnatherum nelsonii (21)

Table 4. Kolmogorov-Smirnov two sample test results. Maximum differences for pairs of variables. G1 L, length first glume;
G2L, length second glume; FL, floret length; FW, floret width; AWN, awn length, ns, differences not significant; *, differences
significant p<.05 >.01; ** p<.01 >.001; *** p<.001.

61L G2L FL FW

G2L QA88^^

FL 0.500*** 0.338***

FW 0-262**, 0-275**^ 0.463***

AWN 0.325*** 0.350*** 0.637*** 0.250*

the length of the lemma, is part of the integrated growth which leads to both the awn and the lemma. The
first thing to initiate will become the awn, the tissue that will become the lemma appears after the awn
when the awn-lemma primordium begins to spread around the floret apical meristem to form the lemma
(Kam & Maze 1974 and references therein). Another developmental feature captured in floret length is
the callus. This is marked by a unique, and often extensive, pattern of cell enlargement slightly oblique to
the longitudinal axis of the floret, at the base of the floret leading to a projection (Maze et al. 1971, 1972;
Mehlenbacher 1970; Kam 1974; Kam & Maze 1974) within which the cells are heavily sclerified. Cells of
the lemma, especially in A. hendersonii and A. wallowaense and to some extent in A. lemmonii are also scleri-
fied. Even though that feature was not measured here, it offers another demonstration of the developmental
complexity of the floret.

Awn length, too, is developmentally complex; the awn is the first thing initiated in the formation of the
floret and its differentiation from the lemma occurs later in development. Its growth in length is a combina-
tion of cell division, apically early in its development and sub-apically later, and cell enlargement, which
appears first in apical cells. Growth in length is limited leading to the shorter awns in A. hendersonii and A.
wallowaense- growth in awn length is greater in A. ocddentale, A. nelsonii and A. lemmonii. Another contribu-
tor to the developmental complexity of the awns of the Stipeae is the sclerenchyma with eccentric lumens
that surrounds the vein in the awn (Maze 1972). This tissue is implicated in the twisting and straightening
of the awn of the Stipeae with hydroscopic changes (Murbach 1900) and is much better developed in A.
lemmonii, A. ocddentale and A. nelsonii. As a further indication of the developmental intricacy in the awn
of the Stipeae, in those awns with well developed sclerenchyma it starts to differentiate much earlier than
surrounding tissues (Maze et al. 1971).

Floret width, also, is developmentally complex since it is the result of two developmental events, the
spread of the awn-lemma primordium around the floret apical meristem followed by subsequent marginal
growth in the lemma. That marginal growth is made more complex through the thickness of the lemma, the
result of periclinal divisions in what could be called the flank meristem of the developing lemma margins.

A strong correlation between allometric coefficients, representing ontogeny, and a phylogenetic
classification, representing evolutionary history, argues for a relationship between the two phenomena. It is

tempting to resurrect the idea of a causal relationship between ontogeny and phylogeny (see Gilbert 2003
& Lovejoy 1959 for a history of such ideas). However, there is a problem with such an argument, the con-
ceptual and empirical gap between the idea of a causal relationship between ontogeny and phylogeny and
the currently popular mechanism proposed for evolution, natural selection favoring certain non-directed
(often called random) variants, is large. And the idea of ontogeny driving phylogeny does not enjoy a well-
established mechanism, even in most modern evo-devo studies that stress how those changes occur or the
description of those changes.

But, such a disconnect is not a part of all modern evo-devo studies. Muller and Newman (2005b) and
West-Eberhard (2005) argue for environmentally induced developmental events as the origin of novelty.
There are a couple of interesting points from their argument. First, Darwin (1859) posited at the origin of
at least some variation as from the conditions of existence, i.e., the environment, as did Lamarck (1809).
Second, the ideas of West-Eberhard (2005) and Muller and Newman (2005b) would seem to be an expres-
sion of Waddington’s (1953) genetic assimilation or the Baldwin effect, i.e., the incorporation of plastic
traits into DNA. Recently Pigliucci and Murren (2003) argued in favor of the Baldwin effect as a source of
evolutionary change. Pigliucci (2007) has also argued for an extended evolutionary synthesis that includes
such phenomena as phenotypic plasticity and epigenetic inheritance, both which have a developmental ba-
sis. And Jablonka and Lamb (1995) and Steele et al. (1998) have described putative molecular mechanisms
whereby environmentally induced traits can be incorporated into the DNA. Other molecular mechanisms
involve methylation of DNA, as well as other chemicals such as ethyl, acetyl and phosphoryl modifications
of histones (Pray 2004).

Another potentially causal relationship between ontogeny and phylogeny has been presented by Maze
et al. (2005), a view derived from the argument that species are information systems (see Brooks 2001, 2002,
2010; Brooks & Wiley 1988; Maze et al. 2005). But there is a depth to the Brooks view not captured in the
modern epigenetic studies or the views of Pigliuccci cited above. Those studies stressed the here and now as
expressed in the material existence of individuals while Brooks arguments see the material existence of the
individual as representing only part of the potential information available to an individual. That potential
information, 1 would argue, represents the information system of the species, an information system that
has captured the history of the species and carries that history forward into the future.

1 find it useful to envision the information system of a species as a code, analogous to the code in a
computer, that captures all the various ways in which information is expressed in the individuals of that spe-
cies. Like the code in a computer, the information system of a species is known to exist when there appears
a specific response in the material world to a certain action. That action, in a computer, could be striking a
key; in a species that action could be the events that stimulate and allow the production of an individual.

Information expression in a species is the result of events mediated by DNA in response to environmen-
tal stimuli, both internal and external. For example, all events, molecular, cytological, histological, leading
to a periclinal division in the protoderm at the apex of a grass floret would become part of the information
system of that species. The same would apply to all other similar events which occur as that plant develops.
The argument that environmentally mediated ontogenetic changes contribute to an expanding informa-
tion system can be seen as part of Darwin’s condition of existence contributing to evolution (Brooks 2010).
Natural selection, which emerges from the interaction of Darwin’s nature of the organism and nature of the
conditions (Brooks 2010), is important as it accounts for survival, a necessary prerequisite for evolution to
occur, i.e., it is necessary but not sufficient for evolution to occur.

As a result of the information expression that accrues through ontogeny of an individual, the informa-
tion system of the species to which that individual belongs would expand. Much of this expansion could be
traced to variation in both internal and external environments that elicit slightly different responses from
the cytoplasm which will, in turn, prompt a different response from the genome. The information system of
a species would also expand as the result of genetic events, viz. mutation, chromosomal rearrangements and
the recombination that accompanies sexual reproduction. As the information system of the species expands

Journal of the Botanical Research Institute of Texas 4(2)

through the appearance of unique developmental events and new arrangements of DNA it becomes unstable
resulting in speciation (Brooks, 2001, 2002, 2010; Brooks & Wiley 1988).

This view incorporates a common causal element into ontogeny and phylogeny. This is seen as a two
phase aspect with a direct phase affecting development of an individual and an indirect phase affecting
evolution. In development the direct cause of the expansion of the information system of a species is the
transformation of matter, the production of information. This production of information accompanies the
energy dynamics of a developing organism; the transformation of matter is the means whereby energy is
processed. I note in passing that it has been shown that an increase in the amount of energy under which
grape leaves develop produces an increase in the diversity of allometric coefficients which also occurs with
both ontogeny and phylogeny (Maze et al. 2003a).

The relation of information expression to energy dynamics in the ontogeny of an individual, the outcome
of the second law of thermodynamics in a highly organized system, offers an indirect tie between energy
dynamics and evolution. The increase in complexity of the information system of a species, an increase
leading to speciation, is indirectly the result of energy dynamics that are the cause of ontogeny. This is not
to say that events such as mutation and recombination do not contribute to the increase in the complexity
of the information system of a species; they do and perhaps may be viewed themselves as a thermodynamic
phenomenon, the increase in informational entropy with the appearance of new things. These ideas are
the same as those arguments first presented by Brooks & Wiley (1988). However, these views do offer a
common cause for ontogeny and phylogeny and, as argued by Maze et al. (2005), such views can offer an
explanation, albeit a controversial one, for incipient speciation that occurs over geographic areas greater
than those occupied by single populations.

ACKNOWLEDGMENTS

Jeanette Whitton performed the phylogenetic analysis used here, Quentin Cronk recommended the use of
coded variables to represent a classification, Cy Finnegan offered unique and valuable insights into the argu-
ments presented here and Dan Brooks, Kali Robson, Edwina Taborsky and Ed Wiley as well as offering insight
into some of the subtleties of phylogenetic analysis. This paper is the result of a long, fruitful collaboration
with Mishtu Banerjee, Kali Robson, Rob Scagel and Peter Sibbald. Their contribution is beyond measure.

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THOMAS WALTER’S SPECIES OE HEDYSARUM (LEGUMINOSAE)

Daniel B. Ward

Department of Botany
University of Florida
Gainesville, Florida 32611, USA.

ABSTRACT

Thomas Walter, pioneer botanist and rice-plantation owner on the Santee River, Berkeley County, South
Carolina, recognized 1056 species in his Flora Caroliniana (1788), the first treatment of American plants to
use the Linnaean sexual system of classification and binomial nomenclature. Of these, over 400 were of
species he believed to be new, while others were taken from the works of Carl Linnaeus. Many of Walter’s
names are in common use today and are readily recognized in American floras by the author designation
“Walt.” But an appreciable number of his names were described so briefiy (in Latin) or without clear distinc-
tion from other species that later authors have been unable to interpret his meaning, either to acknowledge
his names as new or to assign them to appropriate synonymic status.

No index has yet been prepared that fully identifies Walter's names. Later authors have from time to
time dipped into his Flora and determined as best they could the meaning of the names he gave to members
of a given genus or family. Most importantly, Hitchcock (1905) has surveyed the grasses, Dayton (1952) the
pines, Wilbur (2002) the oaks, and Ward (2010) the plants referred to by Walter as Melanthium (Liliaceae). The
present task is to puzzle out the twelve species Walter assigned to the genus Hedysarum (Leguminosae).

Hedysarum L. is not a generic name familiar to botanists of the American Southeast. It was used by Lin-
naeus (1753, 1759, 1762) as a sizable assembly of vaguely related legumes, long since divided into smaller,
more discrete generic groupings. Hedysarum s.s. is still recognized as a genus of about 100 north temperate
species (Mabberley 1997: 331), none of which occur south of Maine or Vermont. The segregate known as
Desmodium Desv. is a rather large, mostly herbaceous genus with peculiar jointed uncinate fruits, while
Lespedeza Michx. is a distinct genus of herbs or sub-shrubs with indehiscent one-seeded fruits and pinnate
trifoliolate leaves.

Because of their commercial importance, the legumes have received thorough taxonomic examination.
Two surveys of the family are of note: the meticulously described legumes of North Carolina (Wilbur 1963),
and the comprehensive understanding of the southeastern U.S. legumes (Isely 1990).

Eastern species of Desmodium fall into two sections: a group of three species with peculiar long-stipitate
fruits well monographed (with their Asian allies) by Isely (1951), and a larger group of species best understood
by Schubert (1950) and supplemented by Isely (1983). American species of Lespedeza have been carefully
studied by Clewell (1966). Indicative of the neglect vested upon many Walter names, none of these authors
has addressed those species of interest hidden in Walter’s Hedysarum.

Association of the names of Hedysarum used by Walter with the names recognized by modem botanists

Journal of the Botanical Research Institute of Texas 4(2)

is not straightforward. Unlike in Quercus, where Wilbur (2002) was equating the dozen Walter names with
a modern-day Berkeley County species-list of near-equal numbers, Walter’s Hedysarum, consisting of both
Desmodium and Lespedeza, is more lengthy and uncertain. Walter recognized twelve species of Hedysarum.
The county-record maps prepared by Wilbur (in Radford et al. 1968) indicate nineteen species of Desmodium
and twelve species of Lespedeza to be expected on the Carolina coastal plain, with still others found not far
beyond. Thus, fewer than half of the species that Walter possibly might have known can be represented in
his Flora.

It is conceivable that the range of species available to Walter was appreciably greater than those found
in the modern coastal plain flora. Though Walter, in the introduction to his Flora, stated he had made his
observations within a 50-mile radius of his Santee River plantation, it has long been recognized that John
Fraser, who in 1787 traveled into the Appalachians and as far south as central Georgia, provided Walter with
materials of otherwise unknown species. Certainly, the Fraser Fir (Abies fraseri, Walter’s “Pinus Cedrus”),
the Showy Ladyslipper (Cypripedium reginae), and other distinctive species could only have come to Walter
via Fraser. The role of Fraser as a source of Walter’s materials should not be overemphasized, however. For
commonplace-appearing species that also occur in abundance on the coastal plain, there is little logic in
Fraser having gathered plants unappealing for horticulture, nor Walter in choosing them for description
over materials available near his home.

Little information is available from herbarium materials. Walter himself kept no herbarium (Ward
2007a). Fraser’s abundant collections were briefly available to Walter, who annotated many of the often-
fragmentary specimens (Ward 2006). These specimens (the Fraser/Walter folio herbarium. Natural History
Museum, London) at times give a clue to the species Walter had described in his Flora, though too often
Fraser’s specimen (perhaps collected in the Carolina mountains) is not the same as the plant intended by
Walter (from near his home on the Carolina coastal plain).

Five specimens of the genus Hedysarum were mounted on a single page (p. 55) of the FraserAValter
folio herbarium (either by Fraser himself or his sons), following Fraser’s return to London in early 1788.
All have now been given page-number/specimen-letter designations (Ward 2006). All five bear three-digit
numbers identified as Fraser’s field numbers, and all five have labels with handwriting identifled as Walter’s.
These specimens, with Fraser’s number, Walter’s label, and its modern identification, are: (1) 55-B: “228”;
“Hedysarum violaceum”; Lespedeza repens (L.) Barton. (2) 55-C; “500”; “Hedysarum Flore magnus”; Desmodium
cuspidatum (Muhl. ex Willd.) Loudon. (3) 55-D: “457”; “Hedysarum”; (crumpled, unidentified). (4) 55-E:
“615”; “Hedysarum”; probably Lespedeza virginica (L.) Britt. (5) 55-F: “721”; “Hedysarum”; Lespedeza capitata
Michx. Only one of these specimens, that of D. cuspidatum (55-C), has been identified as corresponding
to one of Walter’s species of Hedysarum (Fernald and Schubert 1948). A second specimen, that of L. repens
(55-B), bears an epithet used by Walter although his description more closely matches D. lineatum DC. True
L. repens, a common Carolina species, does appear to have been known by Walter, but was misnamed by
him as H. violaceum [= L. violacea (L.) Pers.].

Thus, identification of Walter’s names must be based upon little more than the brief Latin phrase
(often taken from Linnaeus) he used for description, and an estimate of the probability that Walter would
have encountered the plant near his home. Where two species are of similar frequency, yet also similar in
appearance and thus likely not distinguished by him, no single-species identification is possible. Where a
species is common and thus surely known to Walter, the assumption is made that it must be found among
his described species. Rare species are mostly disregarded; only where a species may be rare but with a
distinctive feature seemingly described by Walter, is its rarity as an excluding trait set aside.

The following identifications are not set in stone. Thoughtful consideration of the available data and the
conditions under which Walter worked may cause others to see connections that have been misunderstood
here. Yet even the conclusions drawn here, imperfect as they may be, may have value in giving tentative
meaning to scientific names that for too long have remained obscure.

The species Walter recognized in the genus Hedysarum are listed below, in the sequence as originally

published. An occasional word or phrase italicized and thus emphasized in Walter’s descriptions is shown
here in Roman. Abundance within South Carolina is largely inferred from the county-record maps prep
by Radford et al. (1968). A measure of the likelihood of correct identification is attempted by uniform
age of modifying adverbs preceding the name, where a simple assertion means “little room for doubt,”
“probably” and “possibly” indicate increasing levels of uncertainty.

LIST OF SPECIES

Walter’s name: Hedysarum junceum (p. 184). Linnaeus, Sp. Pi. ed. 2. 1053. 1762; misapplied.

Walter’s description: foliis ternatis lanceolatis, leguminibus uniarticulatis, pedunculis lateralibus subum-
belliferis.

Modern name: Desmodium strictum (Pursh) DC., or Desmodium tenuifolium Torr. <Sr A. Gray
Comments: Both Desmodium strictum and D. tenuifolium are frequent in eastern South Carolina. Walter’s
description is taken directly from Linnaeus, only contracted slightly by deletion of Linnaeus’ “rhombeis” fol-
lowing “uniarticulatis." Not Lespedezajuncea (L.) Pers., of Asia. The often single-loment fruits and lanceolate
leaflets are distinctive of D. strictum and D. tenuifolium, though the description is inadequate to determine
which. No specimen of either of these species is present in the FraserAValter herbarium (BM).

The herbarium does hold a specimen collected by Fraser (55-A), labeled “an Hedysarum” by Walter and
annotated “Hedysarum junceum Walt.” by A. M. Vail. It appears to be Psoralea pedunculata (Mill.) Vail [= P.
psoralioides (Walt.) Cory]. But since Walter clearly intended P. psoralioides to be represented by his “Trifolium
psoralioides” (p. 184), there is no reason to believe the Fraser specimen is also his “Hedysarum Junceum."

Walter’s name: Hedysarum umbellatum (p. 184). Linnaeus, Sp. PI. 747. 1753; misapplied.

Walter’s description: fol. ternatis pedunculis umbelliferis, caule fruticoso recto.

Modern name: Probably Desmodium glutinosnm (Muhl. ex Willd.) Wood

Comments: Infrequent in eastern South Carolina. Walter’s description was taken directly from Linnaeus, with
only recto ' (erect) added. Not Hedysarum umbellatum L., of India. The “umbelliferis" foliage seems unique to
the false whorls of D. glutinosum; leaves of the related D. pauciflorum (Nutt.) DC. are spaced along the stem.
No specimen is present in the Fraser/Walter herbarium.

Walter’s name: Hedysarum barbatum (p. 184). Linnaeus, Syst. Nat. ed. 10. 2:1170. 1759; misapplied.
Walter’s description: / ol. ternatis. floribus racemosis cemuis, calycibus pilosis, leguminibus biarticulatis.

Modern name: Desmodium ciliare (Muhl. ex Willd.) DC.

Comments: Frequent throughout. Walter’s “biarticulatis" loments and “pilosis" calyces well fit this species.
[This identification appears first to have been made by Woods (2008).] Not Desmodium barbatum (L.) Benth.,
of the American tropics. Walter’s description was taken directly from Linnaeus, only slightly reordered.
Articulated fruit confirms that this is a Desmodium. Desmodium fernaldii Schub., D. glabellum (Michx.) DC.,
and D. perplexum Schub., a complex of poorly defined species frequent in eastern South Carolina, were also
considered; but were rejected (fruits often have more than two segments, calyces are scarcely pubescent).
Desmodium strictum (Pursh) DC. and D. tenuifolium Torr. & A. Gray usually have one or two segments, but
appear best assigned under Walter’s “Hedysarum junceum.” No specimen is present in the FraserAValter
herbarium.

Walter’s name: Hedysarum marilandicum ? (p. 185). Linnaeus, Sp. Pi. 748. 1753.
laevibus.

Modern name: Probably Desmodium laevigatum (Nutt.) DC.

Comments: Infrequent in eastern South Carolina, more common westward. Description is modified from
Linnaeus (foliis ternatis: foliolis subrotundis, caule frutescente ramosissimo, leguminibus articulatis laevibus). The
articulated fruits confirm this to be a Desmodium, although no Carolina fruits of that genus are truly smooth

I? I

Journal of the Botanical Research Institute of Texas 4(2)

only along the sutures, otherwise smooth. Desmodium marilandicum (L.) DC. has smooth ovate leaflets (“ovati
emphasized by Walter), though its fruits are “uncinulate-puberulent” (Wilbur 1963), not smooth, and it
quite rare on the South Carolina coastal plain. No specimen is present in the Fraser/Walter herbarium.

Walter’s name: Hedysarumfrutescens ? (p. 185). Linnaeus, Sp. PI. 748. 1753; misapplied.

frutescente). Lespedeza stuevei fits well in that leaflets are ovate to suborbicular and densely pubescent beneath;
it is also “scarcely” fruticose. The name is misapplied; true L.frutescens (L.) Britt. [= Hedysarumfrutescens
L.; Lespedeza intermedia (Wats.) Britt.] is more delicate, less pubescent, and less common (Clewell 1966).
No specimen has been identified in the Fraser/Walter herbarium. [Specimen 55-F appears to be the closely
related Lespedeza capitata Michx.; its label bears only "Hedysarum” in Walter’s hand and “721” in Fraser’s.]

Walter’s name: Hedysarum grandijlorum Walter (p. 185)

Walter’s description: fol. ternatis ovalibus venosis utrique laevibus; racemis axillaribus erectis, leguminibus
pendulis multiarticulatis, Jloribus majoribus.

Modern name: Desmodium cuspidatnm (Muhl. ex Willd.) Loud.

Comments: Infrequent throughout South Carolina. Not Hedysarum grandiflorum Pallas (1773); Walter’s name
is a later homonym and thus illegitimate. Fernald and Schubert (1948: 203) identified specimen 55-C of the
Fraser/Walter herbarium as H. grandiflorum Walt, and referred to it as “Walter’s TYPE.” The specimen was
labeled “Hedysarum Flore magnus" by Walter, who seems not to have recognized it as his own “Hedysarum
grandiflorum,” as named in his Flora. Having been noted by Fernald and Schubert, their typification has been
corrected to neotype for H. grandiflorum Walt. (Ward 2007b).

Walter’s name: Hedysarum viridiflorum (p. 185). Linnaeus, Sp. Pi. 748. 1753.

Modern name: Probably Desmodium viridiflorum (L.) DC.

Comments: Infrequent in eastern South Carolina, common westward. Walter’s description is modified from
Linnaeus (foliis ternatis acutiusculis, caule erecto, racemis longissimis erectis). The “acutiusculis" (slightly acute)
leaves describes Desmodium viridiflorum, but is scarcely unique. No specimen is present.

Walter’s name: Hedysarum hirtum (p. 185). Linnaeus, Sp. Pi. 748. 1753.

Walter’s description: fol. ternatis ovalibus, caule fruticoso, racemis ovatis, calycibus fructibusque hirsutis monos-
permis.

Modern name: Probably Lespedeza hirta (L.) Hornem.

Comments: Frequent throughout. Description is largely original (Linnaeus: /oliis ternatis ovatis, floribusgemi-
natis, leguminibus nudis venosis monospermis). Walter’s description has no point of conflict with L. hirta; its
fruits are densely pubescent. Desmodium ciliare (Muhl. ex Willd.) DC. was also considered; its fruits are often
single-seeded and its calyces pubescent (but scarcely hirsute). If that species, Walter’s epithet has priority
over that of Muhlenberg ex Willdenow (1803). No specimen is present.

Walter’s name; Hedysarum violaceum (p. 185). Linnaeus, Sp. PL 749. 1753; misapplied.

Walter’s description: fol. ternatis ovatis, floribus geminatis, leguminibus nudis venosis monospermis, caule
flexuoso.

Modern name; Probably Lespedeza repens (L.) Barton

Comments; Common throughout. Walter’s description is taken directly from Linnaeus, with only “caule
flexuoso” added. This fits Lespedeza repens rather well (flowers are solitary or in 2s or 3s; fruits are “venosis”
(conspicuously veined), single-seeded; but stem is scarcely flexuous. The name is misapplied (Reveal and

709

Barrie 1991); Lespedeza violacea (L.) Pers. (= Hedysarum frutescens, misapplied) is very rare in South Carolina
(one county). Specimen 55-B was labeled “Hedysarum violaceum” by Walter; it is surely L. repens.

Walter’s name: Hedysarum repens (p. 185). Linnaeus, Sp. Pi. 749. 1753; misapplied.

Walter’s description: /oI. ternatis obcordatis, caule procumbente, racemis lateralibus.

Modern name: Possibly Destnodium lineatum DC.

Comments: If this species, infrequent in eastern South Carolina. This seems the only procumbent Lespedeza
or Desmodium with leaflets broad enough (often obovate) to be considered “obcordatis." Lespedeza repens was
also considered; it is common throughout, racemes are axillary and stems procumbent, but leaflets are mostly
elliptic, and it had also been suggested as Walter’s Hedysarum violaceum. No specimen is present.

Walter’s name: Hedysarum paniculatum (p. 185). Linnaeus, Sp. Pi. 749. 1753.

Walter’s description: /oI. ternatis lineari-lanceolatis, floribus paniculatis, caule erecto.

Modern name: Desmodium paniculatum (L.) DC.

Comments: Frequent throughout. Walter’s description is taken directly from Linnaeus, with “caule erecto”
added. Linear-lanceolate leaflets fit this species, as do erect much-branched (“paniculatis") inflorescences.
No specimen is present.

Walter’s name: Hedysarum nudijlorum (p. 185). Linnaeus, Sp. Pi. 749. 1753.

Walter’s description: Jol. ternatis, scapo Jlorifero aphyllo paniculato.

Modern name: Desmodium nudiflorum (L.) DC.

Comments: Frequent throughout. Description is modified (Linnaeus: Joliis ternatis, scapo Jlorifero nudo, caule
folioso angulato). The long naked scape of D. nudiflorum is diagnostic. No specimen is present.

ACKNOWLEDGMENTS

The author is grateful to Andre F. Clewell and Robert L. Wilbur for many discussions regarding legumes,
to Janet R. Sullivan and Kanchi Gandhi for their helpful evaluation of an earlier version of this manuscript,
to Wendy Zomlefer for her careful review of the present manuscript, and to Christine M. Housel for her
skilled composition of the Spanish abstract.

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LISTADO FLORISTICO Y ASPECTOS ECOLOGICOS DE LA FAMILIA POACEAE
PARA CHIHUAHUA, DURANGO Y ZACATECAS, MEXICO
Yolanda Herrera Arrieta y Armando Cortes Ortiz

Instituto Politicnico Nacional
aiDIR Unidad Durango- COFAA
Sigma 1 1 9 Fracc 20 de Noviembre II
Durango, Dgo., MEXICO, 34220
yherrera@ipn.mx, c_armando25@hotmail.com

RESUMEN

ABSTRACT

introducciOn

Las gramlneas en cuanto a riqueza de especies es la cuarta familia de plantas vasculares, despues de las
compuestas, las leguminosas y las orquideas, y la mas distribuida en el mundo. Se le encuentra desde los
circulos polares hasta el ecuador, en las cumbres de las montanas y al nivel del mar (Lawrence 1951). En
Mexico ocupan el tercer lugar superando en mimero de especies a las orquideas (Rzedowski 1978).

No obstante, parecen ocupar el primer lugar en cuanto a su importancia economica, ya que pertenecen
a esta familia los cereales como el arroz, trigo, maiz y cana de aziicar, base de la alimentacidn humana. La
avena, el centeno, el sorgo y el bambu, tambien de gran importancia para los humanos por proveer de materia
prima para forraje en la produccibn de ganado. Asi tambien el bambu ha sido utilizado ampliamente (desde

712

laboracion de artesanias
a elaboracion de enseres

las antiguas civilizaciones orientales) para la construccion de casas y mueb
y otros productos. En Mexico las especies que son utilizadas regionalmente

reportan por Mejia-Saules y Davila (1992).

Se estima que existen alrededor de 700 generos y 10,000 especies de gramineas en el mundo (Clayton
& Renvoize 1986), de ellos 204 gtoeros y 1,182 especies se encuentran en Mexico (Davila y col. 2006). 117
gdneros (57%) y 505 especies (43%) se distribuyen en esta region del none del pais. Chihuahua cuenta con
98 generos y 383 especies encontrados y registrados a la facha (Herrera y Peterson 2007; Herrera y Cortes
en proceso), mientras que Durango cuenta con 97 generos y 368 especies (Herrera 2001; Herrera y Pamanes
2006 y Herrera, en revision), y Zacatecas con 91 generos y 284 especies segiin el mas reciente recuento
(Herrera y col. 2010). Es la familia de plantas con mayor importancia ecobgica por su diversidad, por su
caracteristica capacidad de formar suelos y porque la mayoria de sus especies son elementos naturales de
vegetaciones primarias, aun cuando un porcentaje no muy alto de ellas (5 a 8% estimado) han desarrollado
la capacidad de dispersarse por el mundo y establecerse como adventicias en habitats diversos.

Por la importancia que reviste la familia Poaceae, el trabajo nomenclatural ha sido cuantioso y elabo-
rado, asi la primicia en la revision de las gramineas de America ha sido llevada a cabo por agrost6logos del
Institute Smithsoniano, quienes ademas de realizar colectas en todo el continente desde finales del Siglo XIX,
de verificar la identidad de los taxa, de describir y aplicar nombres a nuevas especies, para finalmente editar
la informacion obtenida en importantes obras floristicas y floristico-ecologicas de extensas regiones como
Norteamerica (Hitchcock & Chase 1951), de regiones mas localizadas como “La Nueva Galicia” (MeVaugh
1983) o “Los pastizales de Durango” (Gentry 1957). Asi, se inicia el registro en Catalogos, de los nombres
que recibieron los pastes de este continente, a partir de las expediciones de colectores europeos en la Nueva
Espana. Se encuentra que un gran mimero de taxa recibb para si sobs varies nombres en diferentes tiempos
por diversos autores, mientras que algunos nombres (escasos) fueron aplicados de forma repetida a plantas
diferentes. Ambos cases tuvieron que ser reconocidos como sinonimos del nombre con mayor antiguedad
de registro para cada taxon. Quiza el primer catabgo relevante de

el “Indice de gramineas de America” editado por Chase y Niles (1962), quienes incluyeron
aceptados y sinonimia reconocida para su tiempo.

Las Poaceae presentan una distribucidn ecobgica amplia, aiin cuando son los componentes principales
de los pastizales, se encuentran en el sotobosque de los bosques y matorrales de la Sierra Madre Occidental de
Chihuahua, Durango y parte de Zacatecas (Fig. 1). Hay algunas especies que se encuentran desde el bosque
hasta el pastizal, otras en los matorrales y el pastizal, mientras que otras son especificas de condiciones
ecobgicas particulares. Asi, las especies presentan una distribucion mayormente altitudinal, mientras que
un grupo pequeno requiere de un sustrato especifico para sobrevivir. Ejempb de esto son las especies de
suelos con alias concentraciones de sales y las especies acuaticas y subacuaticas.

Durante la realizacion de proyectos financiados por la CONABIO, se genero una base de datos con
aproximadamente 8000 registros de poaceas colectadas en los estados de Chihuahua, Durango y Zacatecas,
provenientes de la revision y registro de datos tornados de ejemplares de herbario, de colectas realizadas
por diversos colectores durante los ultimos 120 anos y depositados en diferentes herbarios del pais y del
extranjero, mismos que fueron registrados en el sistema Biotica proporcionado por la CONABIO. Los sitios
de colecta georreferenciados senalan datos interesantes de las preferencias ecobgicas, distribucidn geografica
y riqueza de los 117 generos y 505 especies presentes en esta region. Esta lista floristica tiene como propbsito

MATERIALES Y METODOS

Se revisaron aproximadamente 8000 ejemplares de herbario, entre ellos algunos 500 ejemplares colectados.

714

los taxa no determinados; tomando como base las descripciones de la bibliografta citada en los generos y
especies. Los herbarios revisados en Mexico por la primera aurora son: CHAPA, CIIDIR, ENCB, HUAZ, lEB,
MEXU y SLPM, y en Estados Unidos: TAES y US. Se realiz6 el analisis de frecuencias a partir de los dates
contenidos en el “Listado floristico de gramineas de Chihuahua, Durango y Zacatecas, Mexico” (Tahla 1).

La determinacion de la identidad de las especies en los ejemplares revisados se llevb a cabo utilizando
claves y descripciones de diversas obras floristicas para Norteamerica (Hitchcock & Chase 1951), para
Mesoamerica (Davidse y col. 1994), y algunas regiones de Mexico (MeVaugh 1983; Herrera 2001; Herrera
y Pamanes 2006; Herrera y col. 2010).

La actualizacidn de la nomenclatura se realizb con base en dos Catalogos: 1) Catalogo de Gramineas
del Nuevo Mundo Qudziewicz y col. 2000; Peterson y col. 2001; Zuloaga y col. 2003 y Soreng y col. 2003),
editado por J.R. Soreng del Instituto Smithsoniano, catalogo que se encuentra en revision y actualizacion
permanente pudiendo ser consultado via internet en: (http://mobot.mobot.orgAV3T/Search/nwcg.html),
2) Catalogo de las Gramineas de Mexico, (Davila y col. 2006); donde se reportan nombres reconocidos y
sinonimia para los taxa mexicanos. Ademas ha sido enriquecida con publicaciones muy recientes de cambios
en la nomenclatura de algunos taxa (ejemplo Finot y col. 2004; Bess y col. 2006; Zuloaga y col. 2007; Bell
y Columbus 2008; Vald6s-Reyna y col. 2009) y/o adiciones de nuevas especies encontradas (ejemplo Finot
y col. 2004; Peterson y col. 2004; Peterson y col. 2006, Peterson y Columbus 2009), asi como la revision
permanente de ejemplares colectados en estos tres Estados.

RESULTADOS

En el Anexo 1 se encuentran relacionados los 536 taxa: generos, especies, subespecies, variedades y formas,
de la regibn norte-centro de Mexico. En ella se relacionan las comunidades vegetales donde se distribuye
cada taxon, el posible origen, la subfamilia a la que pertenecen y por ultimo el estado (Chihuahua, Durango
o Zacatecas) de donde provienen especlmenes de referencia, (escasos son los casos que se citan sin ejemplar

Algunos resultados del analisis de frecuencias se sintetizan en los graficos que se presentan en el pre-
El porcentaje de especies encontradas en cada uno de los estados de la region norte-centro de Mexico

Distribneion y Riqueza de especies.

En la regibn norte-centro de Mbxico los cuatro ecosistemas vegetales principales son:

Bosques templados (encinos, conlferas)

Selvas bajas o bosques tropicales

Matorrales xerbfilos (rosetbfilo, espinoso y mierbfilo)

De estas comunidades la riqueza de zacates se concentra en los bosques templados y pastizales,
que presentan condiciones medias de temperatura, humedad y altitud (Fig. 3).

Las poaceas presentan una distribucibn ecolbgica amplia ya que aunque s
pales de pastizales, se encuentran numerosas especies en el sotobosque de las
asl como en los diversos tipos de matorrales.

Bosques Templados

Estan representados por los bosques de pino, de encino, o mixtos (Pinus, Quercus, Abies, Pseudotsuga, Cupres-
sus yjuniperus). Se distribuyen en la parte alta de la Sierra Madre Occidental, en elevaciones que van desde
2100 a 3250 m s.n.m. En ellos se encuentra la mayor riqueza florlstica de gramineas de la regibn.

Selva Baja o Bosque Tropical Caducifolio

Esta comunidad vegetal se distribuye en la zona de canones y canadas, y es conocida regionalmente como

Tabia 1. Listsdo Mstico de las gramlneas de Chihuahua, Durango y Zacatecas, Mfeo.

Comunidadesvegetales

Achnatherum eminens
Achnatherum hymenoides
Achnatherum multinode
Achnatherum robustum
Aegopogon cenchroides
Aegopogon tenellus
Agropyron cristatus
Agrostis exarata

Andropogon fastigiatus
Andropogon gerardii
Andropogon glomeratus var. pumilus
Andropogon pringlei

Aristidaadscensionis
Aristida appressa
Aristida arizonica
Aristida curvifolia
Aristida divaricata
Aristida eludens
Aristida gibbosa
Aristida gypsophila f. diffusa

Bosque Selva

Aristida. gypsq)hila f. gypsophilides

Aristida laxa var. laxa
Aristida longespica
Aristida pansa f. contracta
Aristida pansa f. dissita
Aristida pansa f. pansa
Aristida purpurea var. fendkriana
Aristida purpurea var. longiseta
Aristida purpurea var. nealleyi
Aristida purpurea var. perplexa

Antndinella berteroniana
Amndinella hispida
Arundo donax f. donax
Arundo donax f. versicolor

Axonopus mexicanus
Bealia mexicana
Blepharidachne bigelovii

ACAD

1 I!

Comunidadesvegetales

Koeleria pyramidata
Lasiacis nigra
Lasiads procerrima

Leersia hexandra
Leptochloa aquatica
Leptochloa dubia
Leptochloa fusca ssp. fascicularls
Leptochloa fusca ssp. uninervia

Leptochloa panicoides
Leptochloa scabra
Leptochloa viscida
Lolium multiflorum
Lolium perenne

Luziola fluitans
Lycurus phalaroides
Lycurus phleoides
Lycurus setosus
Megathyrsus maximus

Metcalfia mexicana
Microchloa kunthii
Mnesithea granularis

Panicum elephantipes

I

I

I

I

g-

I

s

s

732

Numero de especies registradas en

Fk. 2. Porcentajes de espedes encontradas en cada estado de la region.

% de registros

733

“region de las quebradas”. En las estribaciones occidentales de la Sierra Madre Occidental, a lo largo del
limite entre Durango y Sinaloa y en el canon o Barranca del Cobre (Chihuahua). En un intervalo altitudinal

Se registran las siguientes especies de los 28 generos que forman parte de la vegetacion del bosque
tropical caducifolio: Aristida gibbosa, Arundinella hispida, Arundo donax f. versicolor, Axonopus mexicanus, Bo-
thriochloa ischaemum, Bouteloua eludens, B. erecta, B. media, Cenchrus multiflorus, Dichanthelium oligosanthes,
Enteropogon chlorideus, Eragrostis ciliaris, Eriochloa lemmonii, Festuca breviglumis, Gouinia virgata, Guadua
angustifolia, Hyparrhenia ruja, Lasciacis nigra, L. ruscifolia, Leptochloa fusca ssp. minervia, L. panicea ssp. mu-

Paspalum arsenei, P. conjugatum, P. paniculatum, P. pubijlorum, Pereilema ciliatum, Schizachyrium mexicanum,
Setaria latifolia, Setariopsis latiglumis, Sorghastrum incompletum y Tristachya laxa.

Las especies adventicias encontradas en la vegetacion secundaria son: Bothriochloa ischaemum, Eragrostis
ciliaris y Hyparrhenia ruja.

En esta comunidad es donde se encuentra la menor riqueza de gramineas de la region, porque son en
escarpadas y de dificil acceso.

Pastizales

Son comunidades que se distribuyen en la region de los valles, en la vertiente Este o interior de la Sierra
Madre Occidental. Son zonas con precipitacidn anual entre 300 y 400 mm anuales, en altitudes entre 1800
y 2000 m s.n.m.

En este tipo de vegetacion es donde se encuentra la mayor variacidn de pastos naturales, es decir de
origen primario, aqui se distribuye la mayoria de las especies de los generos: Aristida, Bouteloua, Elionurus,
Eragrostis, Heteropogon, Hilaria, Lycurus y Mnesithea, y algunas especies de los generos: Bothriochloa, Chloris,
Digitaria, Erioneuron, Muhlenbergia, Paspalum, Setaria, Sporobolus, Tridens y Urochloa.

Especies que parecen ser exclusivas del pastizal: Achnatherum curvifolium, Aegilops cylindrica, Aristida
purpurea war. jendleriana, Bouteloua parryi var. gentryi, B. trijida, B. wamockii, Bromus inermis, B.japonicus, B.
marginatus, Distichlis littoralis, Echinochloajaliscana, E. muricata var. microstachya, E. walteri, Eragrostis obtusiflora,
E. superba, Hordeum pusillum, Leptochloa aquatica, L. panicoides, L. viscida, Muhlenbergia asperifolia, Panicum
alatum, P. dichotomiflorum, P. elephantipes, P. plenum, Paspalum crinitum, P. tenellum, Phalaris arundinacea y
Sporobolus jimbriatus. ^ ^ ^ ^ ^ ^ ^

especies presentes son de los generos: Aristida purpurea var. longiseta, A. purpurea var. wrightii, A. temipes var.
gentilis, A. ternipes var. ternipes, Arundinella berteroniana, Bothriochloa hybrida, B. laguroides var. torreyana, B.
palmeri, Bouteloua aristidoides, B. barbata, B. breviseta, B. curtipendula, B. eriopoda, B. reederorum, B. scorpioides,
Cenchrus longispinus, Chaboissaea subbiflora, Digitaria insularis, D. sanguinalis, Distichlis spicata, Eragrostis
intermedia var. oreophila, E. sessilispica, Erioneuron nealleyi, E. pilosum, Leersia hexandra, Leptochloa fusca ssp.
fascicularis, L. scabra, Melica montezumae, M. porteri, Muhlenbergia arenicola, M. elongata, M. villiflora, Pani-
cum hallii, Pappophorum bicolor, P. vaginatum, Phalaris canariensis, Pleuraphis mutica, Scleropogon brevifolius.
Sorghum bicolor, Sporobolus airoides, S. contractus, S. cryptandrus, S.giganteus, S. pyramidatus, S. wrightii. Tragus
berteronianus, Tridens muticus, T. texanus, Tripogon spicatus y Urochloa panicoides.

Las especies Cenchrus longispinus, Distichlis spicata, Muhlenbergia asperifolia, Paspalum crinitum, Scleropogon
brevifolius, Setaria leucopila, Sporobolus airoides, S. cryptandrus y S. pyramidatus son especificas de suelos salinos
de pastizales y matorrales.

Especificas de suelos yesosos son Aristida gypsophila, Bouteloua chasei y Sporobolus coahuilensis; en
suelos alcalinos o yesosos las especies: Bouteloua karwinski, B. ramosa, Chaboissaea decumbens, Ch. ligulata,
Ch. subbiflora, Eragrostis obtusiflora, Muhlenbergia asperifolia, M. pauciflora, M. richardsonis, Pleuraphis mutica,
Sporobolus palmeri, S. pyramidatus, S. wrightii.

734 Journal of the Botanical Research Institute of Texas 4{2)

Matorrales Xerofilos

Los matorrales xerofilos, se encuentran en elevaciones entre 1000 y 1800 m s.n.m., con precipitaciones
pluviales de 200 a 400 mm anuales. El Desierto Chihuahuense, es una comunidad dominada por matorral
desdrtico microfilo y/o matorral desertico rosetofilo; los matorrales mas secos de la region. En general son
las comunidades que reciben menores precipitaciones pluviales (100 a 200 mm anuales).

En los matorrales se encontraron las siguientes especies: Aristida gypsophila, A. pansa, Bothriochloa aka,
B. longipaniculata, Bromus arizonkus, Eragrostis barrelieri, E. glandulosa, E. hirta, Hilaria belangeri, Hordeum
arizonicum, Muhlenbergia richardsonis, Paspalum bootteri, Pennisetum polystachion, Phragmites australis, Poly-
pogon interruptus, Rhipidocladum racemiflorum, Schismus barbatus, Setaria adkaerens, S. villosissima, 5. viridis,
Setariopsis auriculata, Sphenopholis obtusata, Sporobolus coahuilensis y S. palmeri.

Las especies encontradas en el matorral del Desierto Chihuahuense son: Aristida purpurea var. perplexa,
Blepharidachne bigelovii, Bouteloua chasei, B. chihuahuana, B. karwinski, B. ramosa, Bromus frondosus, Cenchrus
ciliaris, C. myosuroides, C. pilosus, Chloris rufescens, Cotteapappophoroides, Digitaria cognata, Eriochloa contracta,
E. punctata, Erioneuron nealleyi, E. pilosum, Hesperostipa neomexicana, Muhlenbergia porteri, Munroa squarrosa,
Panicum alatum, Pleuraphis mutica, Schismus arabicus, Setaria verticillata. Sorghum bicolor, Sporobolus jlexuosus,
S. macrospermus, S. spiciformis, Trichloris crinita, T. pluriflora y Tridens albescens. De la region de dunas (Sa-

Especies favorecidas por las actividades humanas, son las siguientes:

Zacates Cultivados y/o Escapades de Cultivo y Arvenses

Las especies que fueron introducidas como cultivo para alimento humano o forraje son: Avena sativa (avena),
Chloris gayana (zacate rhodes), Cynodon dactylon (zacate Bermuda), Cynodon plectostachyus (estrella africana),
Dactyloctenium aegyptium (zacate egipcio), Hordeum vulgare (cebada), Lolium multijlorum (rye grass). Megath-
yrsus maximus (zacate guinea), Panicum antidotale (panico azul), Phalaris canariensis (alpiste). Sorghum bicolor
(sorgo), S. halepense (zacate Johnson) y Triticum aestivum (trigo).

Las especies cultivadas como ornamentales en jardines son: Arundo donax (carricillo), Cortaderia selloana
(pasto de las pampas), Cenchrus ciliaris (buffel), Pennisetum clandestinum (kikuyo), Phalaris canariensis (alpiste)
y Saccharum qfficinarum (azucar).

Anthoxanthum odoratum, Avena fatua, Bothriochloa ischaemum, Cenchrus brownii, C. echinatus, C. longispinus,
Dactylisglomerata, Eragrostis barrelieri, E. curvula, E. lehmanniana, E. spectabilis, Holcus lanatus, Holcus lanatus,
Hordeum arizonicum, Hjubatum, H. murinum ssp. glaucum, Hyparrhenia rufa, Lolium temulentum, Melinis repens,
Mnesitheagranularis, Panicum antidotale, Pennisetum villosum, Polypogon monspeliensis, P. viridis, Schismus barbatus
y Schizachyrium brevifolium son especies no nativas cuya introduccion en la region es probablemente reciente,
considerando el tiempo que llevo la formacidn natural de los pastizales. Generalmente son originarias de
Africa y Eurasia e introducidas involuntariamente, mezcladas con semillas de praderas cultivadas.

Especies introducidas que se comportan como malezas en terrenos de cultivo o en areas de disturbio en
diversos tipos de vegetacion como: Avena fatua. Cenchrus ciliaris, C. incertus, C. myosuroides. Digitaria ciliaris.
D. filiformis, D. sanguinalis, D. ternata, Eleusine indica, E. multiflora, Eragrostis cilianensis, E. ciliaris, E. maypu-
rensis, Lolium perenne. Setaria adhaerens, S. verticillata, S. viridis. Sorghum halepense y Tragus berteronianus.

Otro grupo de especies importantes son las favorecidas por la presencia de agua:

Especies Acuaticas, Subacnaticas o Tolerantes

Son especies que se encuentran en condiciones humedas hasta llegar
intermitente, ya que las lagunas, canales o escurrimientos en esta reg
del ano. En la Tabla 1 del Anexo se marcan 24 especies acuaticas c
tolerantes con T. De este tipo de microhabitat se encuentran 42 taxa
riparia y 7 mas con BG del bosque de galena.

on VA, 14 subacuaticas
con VR que indica son d

C3 7C,:

736

Origen

Fkl 5. Espedes nativas por su distribudon a nivel continente.

737

Subfamilias

AGRADECIMIENTOS

Se agradece el financiamiento otorgado a los proyectos: “Floristica de Gramineas de Durango” DGPI-1998045
y CONABIO-R035, “Floristica de Gramineas de Zacatecas” SIP-20070429 y CONAB1O-EE014; “Floristica de
Gramineas de Chihuahua” SIP-20100879 y CONABIO-GE003; donde se generaron los datos incluidos en
el presente reporte. Se agradece a la Dra. Teresa Mejia Sauks y un revisor anonimo su tiempo para revisar
y hacer observaciones que enriquecieron el presente trabajo.

REFERENCIAS

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Clade’ (Poaceae: Paniceae). Syst. Bot. 31:656-670.

Chase, M.A. y C.D. Niles (compilers) 1 962. Index to grass species. 3 vols. G.K. Hall, Boston.

Clayton, W.D. y S.A. Renvoze. 1 986. Genera graminum grasses of the World. Kew Bull. Addit. Ser. 1 3. Royal Botanic
Gardens, Kew. ^ „

Davidse, G., M. Sousa S. y A.O. Chater (eds.). 1 994. Flora Mesoamericana Vol. 6: 1 -543. UN AM, Missouri Bot. Gard. &
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MIRANDEA GRISEA (ACANTHACEAE), NEW FOR

COAHUILA AND DURANGO, MEXICO

Eduardo Estrada-Castillon’ Jose Angel Villarreal-Quintanilla

Universidad Autonoma de Nuevo Leon, A.P. 4
Facultad de Ciencias Forestales, 67700
Linares, Nuevo Ledn, MEXICO

Universidad Autdnoma Agraria Antonio Narro
Departamento de Botdnica
2531 5 Buenavista, Saltillo

Jorge Arturo Alba-Avila

Departamento de Botdnica
Escuela Superior de Biologia
Universidad Judrez del estado de Durango
35010, Gomez Palacio, Durango, M&dCO

ABSTRACT

RESUMEN

The genus Mrandea was described by Rzedowski (1959) based on Mirandea grisea from San Luis Potosi. At
present six species are known in the genus, all of them endemic to Mexico: Mirandea huastecensis T.F. Daniel
restricted to the Canyon La Huasteca in the state of Nuevo Leon; M. hyssopus (Nees) T.F. Daniel and M. nutans
(Nees) T.F. Daniel to Queretaro and Hidalgo; M. sylvatica Acosta to Chiapas, Oaxaca, Tabasco and Veracruz;
and M. andradenia T.F. Daniel to Tamaulipas (Daniel 2003; Daniel & Acosta 2003). Mirandea species resemble
those of Carlowrightia, however, they differ in stamen characteristics. In Mirandea the filaments are twisted
and appressed to the upper lip of the corolla, the anthers open to lower lip, and the style is recurved near
the apex, while in Carlowrightia, the stamens are not appressed to the upper lip of the corolla, the filaments
are not twisted, the anthers open to the upper lip, and the style is straight near the apex (Daniel 1982).

Mirandea grisea has been recorded from San Luis Potosi near El Tepeyac (MEXU, TEX), Matehuala
(MICH), Guadalcazar (ENCB, TEX) and El Huizache (ENCB, TEX) (Daniel 1982) and in Tula in southwest-
ern Tamaulipas (CAS, MEXU, MO) (Daniel 1999). Plants occur in scrub vegetation. The new localities are
the following:

Distribution and Conservation.— The new records for this species are in the Sierra of Jimulco (ca. 40 km N of
Torreon in Coahuila), and in the Sierra El Samoso (ca. 10 km W of Sierra de Jimulco, in Durango) (Fig. 1). Veg-
etation of both areas is desert scrub with Larrea tridentata. Agave lecheguilla, Parthenium incanum, P. argentatum.

740 Journal of the Botanical Research Institute of Texas 4(2)

Opuntia lindheimeri, Fouquieria splendensjatropha dioica, and Lippiagraveokns associations. Both localities are
almost 300-320 km NW from the nearest locality were Mirandeagrisea was recorded previously (El Huizache
area). In Jimulco as in El Sarnoso, Mirandea grisea is rare; it does not form extensive stands as in San Luis
Potosi (Daniel 1982). Only a few individuals were located scattered through the vegetation on inclined planes
and in the valleys at 1,500 to 1,700 meters. Canopy cover was less than 30%. The new records for this species
expand its known distribution almost 200 km northwestward in the Chihuahuan Desert Region. Mirandea
grisea could be included into the NOM-059-ECOL-2001 (Semarnat 2001) as endangered species in Coahuila
and Durango States, since there are only few populations and few individuals in each of these populations.

ACKNOWLEDGMENTS

We thank to T.F. Daniel and one anonymous reviewer for their helpful comments and to Barney Lipscomb
for his comments in the manuscript and reviewing the final English version of the manuscript.

REFERENCES

Daniel, T.F. 1 982. The genus Mirandea (Acanthaceae). Contr. Univ. Michigan Herb. 1 5:1 71-1 75.

Daniel, T.F. 1999. Nuevos registros estatales de Acanthaceae en Mexico, Bol. Inst. Bot. (Univ. Guadalajara)
7(1 -3):5 1-59.

Daniel T.F. 2003. A new combination in Mirandea (Acanthaceae). Acta Bot. Mex. 62:9-13.

Daniel, T.F. y S. Acosta-C 2003. Familia Acanthaceae. Flora del Bajio y Regiones Adyacentes. Fasci'culo 1 1 7. Instituto
de Ecologia, A.C. Patzcuaro Michoacan.

Rzedowski, J. 1959. Nuevas adiciones a la flora mexicana. Ciencia (Mexico) 19:80.

Sermarnat. 2001. NOM-059-ECOL-2001 (lista de especies amenazadas, en peligro de extincion y sujetas a pro-
teccion especial). Diario Oficial de la Federacion, Mexico, D.F

REGISTRO DE DOS NUEVAS LOCALIDADES Y REUBICACION DE INDIVIDUOS
DE UNA DE ELLAS DE ASTROPHYTUM MYRIOSTIGMA (CACTACEAE)

EN DURANGO, MEXICO

Jaime Sanchez Salas, Gisela Muro Perez,

Eduardo Estrada Castillon y Mario Garcia Aranda

Facultad de Ciencias Forestales
Universidad Autdnoma de Nuevo Leon
Apartado postal 41, 67700, Linares, N.L, MBdCO

El Desierto Chihuahuense se caracteriza por su rica diversidad y endemismo de cact^ceas (Hernandez y
Godinez 1994; Hernandez y Barcenas 1995), muchas de las cuales se encuentran en alguna categoria de
riesgo (Hernandez y Godinez 1994; Hernandez y BSrcenas 1995 y 1996; Gomez-Hinostrosa y Hernandez
2000; Hernandez et al. 2001). Una de las especies catalogada como amenazada de extincion es Astrophytum
myriostigma Lem. (SEMARANT 2002). Especie distribuida en los estados de Coahuila, Nuevo Leon, Tamau-
lipas, San Luis Potosi (Bravo y Sanchez-Mejorada 1991; Guzman et al. 2003; Henrickson yjohnston 1997) y
Durango, habitando pendientes pedregosas, asociado a comunidades de Agave lecheguilla y Hechtiaglomerata
(Henrickson yjohnston 1997). Actualmente se reconocen dos variedades de esta especie Astrophytum myrios-
tigma var. myriostigma y Astrophytum myriostigma var. coahuilense (Kanfer) Borg, esta ultima con alto grado
de polimorfismo (Henrickson yjohnston 1997), por lo que llega a ser considerada como una sola especie
(Guzman et al. 2003). Es una especie evolutivamente calcicola apomorfica (Del Castillo 1996) y al menos
en las localidades del Desierto Chihuahuense donde se ha registrado, se distribuye con rangos altitudinales
que van de los 1170 a 1205 msnm, con suelos de tipo calcareos (Sanchez-Salas 2002). Esta especie no fue
reportada para el estado de Durango (Guzman et al. 2003).

Poblacion de Astrophytum myriostigma en la Sierra El Sarnoso

En junio de 2002 se registrb una nueva poblacidn de este taxa (Fig. 1) en Durango, localizada en la sierra
“El Sarnoso” (25°35’N, 103°35'W), dentro del municipio Lerdo, Durango (S^nchez-Salas y Romero 2003).
La vegetacion en esta area corresponde a matorral rosetofilo, predominando Agave lechuguilla Torr., Opuntia
leptocaulis DC. y Jatropha dioica Sesst, esta poblacion es diferente a la reportada por Henrickson yjohnston
(1997), localizada el Cerro “La bola” en las montanas cercanas al poblado de Viesca, 350 Km al Noreste de
Torrebn, Coahuila.

La poblacion de A. myriostigma localizada en la sierra “El Sarnoso” abarca una superficie de aproxi-
madamente 7.11 KmL Con la finalidad de conocer la densidad de individuos de la poblacion, en Junio de
2002 realizamos un primer muestreo estratificado (Franco 1985) geoposicionando ocho cuadrantes de 10

742 Journal of the Botanical Research Institute of Texas 4(2)

alyEndna2005

743

X 10 m, donde registramos un total de 75 individuos (Muro-Perez 2002). Posteriormente, en Septiembre del
mismo ano, se geoposicionaron otros ocho cuadrantes en lugares diferentes dentro de la misma poblacion,
esta vez se registraron 27 individuos mas, totalizando 102 individuos (Sanchez 2002; Sanchez y Romero
2003). En Octubre de 2006, utilizando la misma metodologia, se cuantificaron de nuevo los individuos de
A. myrisotigma en los mismosl6 cuadrantes. En esta ocasion el registro de individuos disminuyd a 92 indi-
viduos, 10 individuos menos que en Septiembre de 2002 (registro anterior). Esta disminucion fue resultado
de la extraccion ilegal de las plantas (Muro-Perez 2006), pues se pudo constatar visualmente los huecos en
la tierra que dejan las plantas al ser extraidas.

En Mayo de 2009, se realize otra cuantificacion de individuos de esta especie en los mismos cuadrantes
y con la misma metodologia, en esta ocasion se registraron unicamente 75 individuos en la poblacion, 35
menos que los registrados en Octubre de 2006. Estas 35 plantas saqueadas, fueron localizadas a 40 m de
distancia respecto la poblacibn origen de esta especie, en dos costales, con 15 y 20 individuos respectiva-
mente. Desafortunadamente, los individuos extraidos eran individuos adultos, de 35 cm de largo o mayores,
reportados como “plantas madre” (Reza 2008), mismos que proveen las semillas que mantienen a la poblacion

reemplazaran a los adultos seniles.

La extraccion ilegal de esta especie en la zona es llevada a cabo por pobladores de la region, como
encargo de traficantes extranjeros que pagan cantidades irrisorias por ejemplares de esta especie (S^inchez-
Salas 2002; Muro-Perez 2002). En el presente ano se realizara una nueva cuantificacion de individuos, ya
que de acuerdo con los pobladores del area, esta zona aun no esta exenta de saqueo y comercio ilegal por
parte de coleccionistas extranjeros quienes pretenden poseer esta codiciada planta dadas sus peculiaridades
morfolbgicas de gran belleza.

Poblacion de Astrophytum myriostigma en el Ejido Villa Nazareno

El Ejido Villa Nazareno (Fig. 1) se localiza en el Municipio Gomez Palacio, Durango, Mexico (25“24'N,
103°25'W), la vegetacidn en esta zona corresponde a comunidades de matorral rosetofilo, donde las especies
predominantes son; Agave lecheguilla Torr, Yucca torreyi Shafer, Larrea tridentata Coville y Hechtia glomerata
Zucc. En febrero de 2004 registramos la presencia de 3 individuos de esta especie, la poblacion se delimito
geograficamente con geoposicionador para conocer la superficie de distribucion, la cual abarca aproximada-
mente 25 m^. En octubre de 2004, se geoposicionaron 6 cuadrantes de 10 x 10 m dentro de esta poblacion
para cuantificar el numero de individuos, en total se cuantificaron 32 plantas de A. myriostigma. Adyacente a
esta poblacion de individuos, se localiza un area donde actualmente se extrae mineral (caliche) y su Area de
impacto se expande hacia esta poblacidn de A. myriostigma. Una semana despues, localizamos otra pequena
poblacion de individuos a una distancia de aproximadamente 50 m al E de la primera poblacion y al igual
que la poblacion anterior, la delimitamos geograficamente con GPS. Esta abarca una superficie de 250 m^
donde se establecieron tres cuadrantes de 10 x 10 m para calcular la densidad de individuos en su superficie.
En esta poblacion se registraron 10 individuos, en total ambas poblaciones suman 42 individuos.

Para evitar la destruccion de la poblacihn y muerte de los individuos de la poblacion adyacente al area
de extraccion de mineral de caliche, se presento un programa de rescate de esta especie a SEMARNAT (Se-

la segunda de estas; concedida la autorizacion (No. Oficio; SG/1 30. 2. 2. 1/002 193; No. Tramite; 3532), reubi-
camos los 32 individuos a la segunda poblacidn. En la actualidad, los individuos parece se han adaptado bien
al sitio donde se reubicaron, pues no ha habido mortandad en ninguno de ellos; seguiremos monitoreando
esta poblacion para tratar de seguir protegiendo esta especie en inminente peligro de extincidn.

La presencia de las dos poblaciones de A. myriostigma registradas para el municipio de Gdmez Palacio
y Nazareno, Dgo., con una separacidn entre ellas de 37 km lineales; aunada a la del Cerro Bola cerca de
San Pedro de las Colonias (Britton y Rose 1963; Henrickson y Johnston 1997; Villarreal y Encina 2005) en
Coahuila (Fig. 1); separadas por 50 km lineales en promedio, respecto a los nuevos registros (Fig. 1); hacen
suponer preliminarmente que estas poblaciones se encuentran sujetas a la dispersidn llamada “area de Noe”

Sanchez etal.J

Muro-P^rez, G. 2006. Efecto de la altitud y las propiedades del suelo sobre la densidad poblacional deAstrophytum
myriostigma.Jes\s de maestria. InstitutoTecnologico deTorreon (ITT). Pp. 72-74.

Muro-PErez, G., U. Romero-MEndez, J.D. Flores R. y J. SAnchez-Salas. 2009. Algunos aspectos sobre el nodrizaje en Asfro-
phytum myriostigma Lem. (1839) (Cactaceae), en la sierra El Sarnoso, Durango, Mexico. Nakari 20(3):43-48.

Reza-Carrillo, M. 2008. Aspectos demograficos de una poblacion de Astrophytum myriostigma Lemaire (1839)
Cactaceae en la sierra el Sarnoso, Durango, Mexico. Tesis de Licenciatura. Escuela Superior de Biologia. Uni-
versidad Juarez del Estado de Durango. P. 37.

Rzedowski, J. 1 978. Vegetacion de Mexico. Limusa. Mexico, D.F.

Sanchez-Salas, j. y U. Romero-MEndez. 2003. Listado de la flora cactoldgica de la Sierra El Sarnoso, Durango, Mexico.
Nakari 14:27-36.

SAnchez-Salas, j. 2002. Distribucion geografica de la flora cactologica con estatus de conservacion, de la sierra
el Sarnoso, Durango, M^x. Tesis de licenciatura. Universidad Juarez del Estado de Durango. Escuela Superior
de Biologia. P. 13.

Sanchez-Salas, j., G.E. MarKnez, y RJ. Flores. 2006. Efecto del tamano de semillas en la germinacion deAstrophytum
myriostigma Lemaire (Cactaceae), especie amenazada de extincion. Interciencia 31(5);371

SEMARNAT. 2002. NORMA Oficial Mexicana NOM-059-ECOL-2001. Proteccion ambiental-Especies nativas de
Mexico de flora y fauna silvestres-Categorias de riesgo y especificaciones para su inclusion, exclusion o
cambio-Lista de especies en riesgo. Secretaria de Medio Ambiente y Recursos Naturales. Diario Oficial de la
Federacion. Mexico, D.F.

Villarreal-Quintanilla, A.J. y AJ. Encina-DomInguez. 2005. Plantas vasculares endemicas de Coahuila y Algunas Areas
Adyacentes. Acta Bot. Mex. 70:1-46.

746

BOOK REVIEW

Eugene S. Hunn. 2008. A Zapotec Natural History: Trees, Herbs, and Flowers, Birds, Beasts, and Bugs
in the Life of San Juan Gbee. (ISBN 978-0-816-52617-8, hbk.). University of Arizona Press, 355
South Euclid Avenue, Suite 103, Tucson, Arizona 85719, U.S.A., (Orders: Avww.uapress.edu, 1-800-
426-3797). $50.00, 262pp., CD-Rom, 11 b&w photos, 6" x 9".

ERIGERON MANCUS (ASTERACEAE) DENSITY AS A BASELINE TO DETECT
FUTURE CLIMATE CHANGE IN LA SAL MOUNTAIN HABITATS

James F. Fowler

Barb Smith

USFS Rocky Mountain Research Station
Flagstaff, Arizona 8600 1 USA
jffowler@fs.fed.us

ABSTRACT

748

Service 1994). Forest Service objectives for these areas include protection against serious environmental
disruptions and serve as baseline areas for measuring long-term ecological changes. No permanent study
plots were established in the Mt. Peak RNA prior to our climate change work on E. mancus.

Global mean temperature is predicted to rise 1-6° C in the next century due to increased concentration
of greenhouse gases (Solomon et al. 2007) and has increased 0.4° C over the previous 150 years (Trenberth et
al. 2007). The predicted general response of species to this increased warmth is to migrate north in latitude
or up in altitude (Grabherr et al 1994; Theurillat & Guisan 2001; Walther 2002; Root et al. 2003; Kullman
2008; Lenoir et al. 2008) although most ecologists expect species to respond individualistically and not as
intact communities (Franklin et al. 1991; Theurillat & Guisan 2001). Alpine species population decline and
extinction are also possible since there may be little available habitat for species’ upward migration (Chapin
& Korner 1994; Grabherr et al 1994; Theurillat & Guisan 2001). These potential ecological changes indi-
cate the need to establish baseline plant species’ distributions and abundances at local scales to definitively
detect changes (Post et al. 2009). Locally abundant, single mountain endemic species like E. mancus offer
good opportunities to establish baseline studies for this purpose.

Our primary goal in this study was to measure basic population biology parameters for the E. mancus
population on the ridge from Mt. Laurel in the Middle Group of the La Sals west to treeline. 'We estimated
plant density and patch size in order to estimate the total number of E. mancus plants on this ridge. A sec-
ondary goal was to describe vascular plant species composition within the area populated by E. mancus. For
both of these goals, we were also interested in the influence of elevation within alpine habitats. We addressed
these research questions in the context of current and predicted global warming and the need to establish
baseline ecological information in order to understand future climate change effects.

METHODS

The study area was in the Middle Group of the La Sal Mountains in San Juan County in southeastern Utah
(Fig. 1) on the Manti-La Sal National Forest. The study area was defined as the Mt. Peak Research Natural
Area (RNA) and the ridge just west of Mt. Laurel. In June, 2009 we established a 1-km ekvational ridgeline
transect from timberline to the large talus field at the west base of Mt. Laurel. This included three vegetation
patches with gaps for the large talus patches near the USFS pre-Laurel weather station. It covers an ekvational
range from 3430 m to 3629 m through patches of alpine herbaceous vegetation. We measured both E. mancus

July near peak flowering time. Vascular plant species composition was measured at 20-m intervals along
the above transect with a random start sampling location within first 20 m and at systematic 20-m intervals
thereafter. Erigeron mancus density was measured at randomly chosen points along E. mancus patch widths
at the same 20-m intervals along this transect. In August 2009, we established a 100-m long E. mancus den-
sity transect along the Middle Group of the La Sals crest line in the saddle just south of Mt. Laurel. Density
measurements were taken as above using this transect as a baseline to measure patch widths. Ekvational
range of this transect was 3632-3642 m. We recorded latitude, longitude, and elevation at each sampling
frame with Trimble* Geo XT 2005 Series GPS at sub-meter accuracy using the North American Datum 1983.

Voucher specimens of vascular plant species were collected in June, July, and August. Plants were identi-
fied using descriptions and keys published in FNA (1993+) and by comparison with known specimens in the
Rocky Mountain Herbarium in Laramie, WY and the USFS Herbarium at RMRS in Flagstaff, AZ. Specimens
are deposited at the latter herbarium. Plant nomenclature follows FNA (1993+) and the Intermountain Flora
(Cronquist 1972+) in that order of priority. Descriptive statistics for plant densities and species centroid
elevations were calculated with SAS/STAT 9.2 (SAS 2008).

RESULTS

Erigeron mancus was confined to dry ridgelines along the ekvational transect (Fig. 2). It was not found in
large, loose talus areas and tended to sharply decrease in abundance near more mesic areas, especially where

jLaSal Mountains!

snow appeared to persist later into the growing season. Plant counts per sampling frame ranged from 0 tc
35, reflecting the species’ visual patchiness. A range of plant sizes was observed with the smaller ones hav-
ing a single unbranched caudex and the larger ones having multiple caudex branches. We did not measure
plant size or age but some appeared to be relatively young with a small diameter at the top of the caudex
while others appeared to be much older with a relatively large diameter caudex and/or a pedicellate caudex
due to soil erosion. Mean density was 7.09 plants/m^ (Table 1) which yielded a population estimate of over
200,000 plants along Mt. Laurel ridge and its nearby southern Crestline. Density does not appear to change
significantly with elevation since the standard errors of the density estimates of the three main patches
overlap (Table 1). The largest E. mancus patch size with the largest number of plants is located above and
just east of the USFS pre-Laurel weather station (Table 1, Fig. 2).

The elevation of the sampled E. mancus population centroid weighted by E. mancus density was 3537
m (12,330 ft) which is within the largest patch near a shallow windswept saddle east of the weather station
(Fig. 2). We also found a small outlier patch in an open area well within the spruce-fir forest at 3356 m
(11,010 ft) and 74 m below the next patch at the timberline start of our sampling transect at 3430 m (11,247
ft). There are additional, unsampled patches of E. mancus along the Crestline of the Middle Group of the La
Sals and at the north base of Mt. Mellenthin.

Vascular plant diversity along the Mt. Laurel ridge transects averaged 17 ± 0.58 SE species per square me-
ter with a richness range of 10-26 species per square meter. We collected one new Utah state record, Artemisia

752

Journal of the Botanical Research Institute of Texas 4(2)

For the 38 species encountered along the elevation transect five or more times, the elevation of population
centroids were calculated based on occurrence within individual sampling frames then placed in ascending
elevation order (Table 2). Most species in the middle part of Table 2 occurred fairly often and ranged over
most of the transect’s elevation range, 3430-3629 m. Gentiana parryi, Draba abajoensis, (frigeron grandijlora,
Carex rossii, and Calamagrostis purpurea were restricted to the lower part of the elevation range, <3550 m.
Silene acaulis. Trifolium nanum, Androsace chamaejasme, Minuartia obtusiloba, and Poaglauca subsp. glauca were
restricted to the upper part of the elevation range, >3481 m. Two species, Draba aurea and Elymus scribneri,

tively narrow standard errors. The two varieties of Potentilla ovina had well separated centroid elevations
with non-overlapping standard errors indicating that those centroid elevations are significantly different.
However, that is not the case between Poa glauca subsp. glauca and Poa glauca subsp. rupicola where the
centroid standard errors overlap, thus indicating no significant difference.

DISCUSSION

Prior to this study little was known about the species density of E. mancus, however the population along
the dominant plant species as well as other meadow areas with dense forb/graminoid cover in which E.

between the major patches we measured (Table 1) and the observed range of plant sizes and presumable
ages would support the hypothesis of a stable population. Similarly, its range from timberline to crest line,
including the additional population patches we documented last year (Smith 2008), indicate that it may
be quite widespread within the Middle Group of the La Sals. Thus £. mancus seems to be persisting under
current levels of anthropogenic activity and the current climate pattern. Whether this will remain so under
a warming climate is a much more open question.

The population centroid elevation, frequency of occurrence, and elevation range data for the species
associated with E. mancus shown in Table 2 provide the baseline data for future comparisons. Significant
changes in these measures may represent ecological change due to climatic or anthropogenic influences. The
elevational data for each species along this transect represent their ecological amplitude along this elevational
gradient using raw elevation as a surrogate for temperature, wind, precipitation, and other variables that
describe the ecological niche of each species. We now have precise spatial coordinates for multiple occur-
rences of 38 vascular plant species along this elevational transect which should make detection of impending
local extinction possible.

The main rationale for Candidate 2 status species is that sufficient data on biological vulnerability and
threat are not currently available to propose listing a species as Threatened or Endangered and that Candi-
date 2 listing will encourage the necessary research (U.S. Fish and Wildlife Service 1993). They also listed
status trend as Unknown for E. mancus to indicate that additional survey work is required to determine
current trends. This study provides current baseline data on a few aspects of the population biology of E.
mancus which will then allow future re -measurements of density, population size, and elevational centroids
to indicate population trends in response to climate change and anthropogenic stressors.

ACKNOWLEDGMENTS

Thanks to Addie Hite and Brian Casavant for help with fieldwork and to the Canyonlands Natural History
Association, Moab, Utah for funding this research. The final report for this research is posted on the Can-
yonlands Natural History Association website: http://www.cnha.org. We also thank the Rocky Mountain
Herbarium for assistance in plant identification. Richard D. Noyes and Timothy K. Lowrey provided excellent

SCALLOPLEAF SAGE (SALVIA VASEYl: LAMIACEAE) DISCOVERED IN ARIZONA

James W. Cain, IIP

Richard S. Felger

Jniversity of Arizona Herbarium
Herring Hall
University of Arizona
Tucson, Arizona 8572 U.S.A.

Brian D. Jansen^

School of Natural Resources
University of Arizona
Tucson, Arizona 85721 U.S.A.

Paul R. Krausman^

School of Natural Resources
University of Arizona
Tucson, Arizona 85721 US. A.

ABSTRACT

INTRODUCTION

On 11 February 2003, JWC and BDJ were collecting and measuring vegetation near the crest (ca. 840 m)
of the Sierra Pinta (113.56051°W, 32.29510°N, NAD 83) on the Cabeza Prieta National Wildlife Refuge in
southwestern Arizona. These efforts were part of a larger study investigating the ecology of desert bighorn
sheep (Ovis canadensis mexicana). They encountered a highly aromatic shrub that they could not identify and
collected a dried inflorescence and portions of the herbage to be identified by botanists at the University of
Arizona Herbarium. The plant was identified by RSF as scallopleaf sage, Salvia vaseyi (Porter) Parish (Audibertia
vaseyi Porter), and represented a new record in Arizona. Cain subsequently collected additional specimens,
and in 2010 J. Malusa (School of Natural Resources, University of Arizona, pers. comm.) discovered another
population in the nearby Copper Mountains in the Goldwater Military Range (see Broyles et al. 2007 for
geographic information). This species was previously known from southeastern California and northernmost
Baja California on the western edge of the Sonoran Desert in the Peninsular and eastern Transverse Ranges
and adjacent mountains (e.g.. Consortium of California Herbaria; Hickman 1993. Fig. 2).

Salvia vaseyi is a highly aromatic shrub or subshrub reaching ca. 1+ m tall (Figs. 1 & 3). It is facultatively
drought deciduous and experiences severe dieback in drought. The leaves are thick and highly variable
in size, depending on soil moisture, densely pubescent with minute, appressed trichomes, conspicuously
glaucous, and the surfaces are often rugulose. The inflorescences are verticillate on usually wand-like, tall
flowering stems (Figs. 1 & 3). The calyces, 8-14 mm long, have mostly awned lobes, the awns becoming

J. Bot. Res. Inst Texas 4(2): 755 - 760. 2

757

Fk. 2. Distribution of 5a/wa vaseyi.
edu/cgi-bin/get_consort.pl) and t
California lotalities based on sped

spinescent when dry after seed maturity. The corollas, filaments, and styles are pure white, and the corolla
tube 13-20 mm long (Figs. 1 & 3). Fruits have light brown nutlets 2.5-3 mm long. The Arizona plants are
reproductive at least in spring and likely in fall, depending on rains (soil moisture). We found no morpho-
logical differences between Arizona and California specimens.

The Arizona S. vaseyi populations occur in vegetation communities characteristic of granitic mountains
in the Lower Colorado Valley phytogeographic region of the Sonoran Desert (Shreve 1951; Turner & Brown
1994; Felger et al. 2007). Common plants growing in association with S. vaseyi include foothill palo verde
(Parkinsonia microphylla), brittlebush (Enceliafarinosa), creosotebush (Larrea divaricata var. tridentata), des-
ert lavender (Hyptis emoryi), white bursage (Ambrosia dumosa), rough jointfir (Ephedra aspera), range ratany
(Krameiiagrayi), desert agave (Agave deserti), ocotillo (Fouquieria splendens), buckhorn cholla (Cylindropuntia
acanthocarpa), barrel cactus (Eerocactus cylindraceus), saguaro (Carnegieagiganted), and elephant tree (Bursera
microphylla).

The climate in Yuma County, Arizona, is extremely arid. Rainfall is bimodal and varies widely. The
Arizona S. vaseyi sites are probably nearly frost-free and probably experience 7.6-10.2 cm (2.86-4.18 in)
average annual rainfall based on the nearest long-term weather data from Wellton and Yuma (Western Re-
gional Climate Center 2010). Mean rainfall recorded in the Sierra Pinta (2002-2005) was 1.7 cm in summer
(April-August) and 4.9 cm in winter (December-March). High temperatures in summer reached >45°C,
and low temperatures in winter reached 2.6°C QWC, unpublished data).

Salvia vaseyi in the Sierra Pinta was found in 3 local areas, each approximately 60 m in diameter and on
steep granitic slopes with southwest exposures. The plants at the summit, at 838 m (2750 ft; Cain & Jansen
11 Feb 2003), were common but not abundant. The second site, on the lower slope of the mountain at ca. 405
m (1180 ft; Cain & Jansen 15 Nov 2003) and 6 km north of the first locality, had 20-30 S. vaseyi plants. The
third site, near the base of the mountain (ca. 360 m; Cain & Jansen 21 Feb 2005) and 5 km northwest of the

first site, had less than 12 individual plants. On 8 March 2010, J. Malusa found another Arizona population
in the Copper Mountains on a north-facing slope about 120 m below the summit. The plants were reported
as common at this site. The Copper Mountains are about 28 km west-northwest of the Sierra Pinta (Fig 2).

During extensive fieldwork over several years, JWC and BDJ did not find other S. vaseyi plants in the
Sierra Pinta or to the west in the ecologically similar Cabeza Prieta Mountains, and J. Malusa reported that
he did not find additional plants in the Copper Mountains or similar nearby mountains. Felger and others
have conducted extensive fieldwork in other granitic mountains in southwestern Arizona and also have
not found other S. mseyi plants. Yet, due to the remote location of many southwestern Arizona mountains
and restricted vehicle access due to wilderness designation, there might be other localities where this or
other species, previously undocumented in Arizona, occur. Salvia vaseyi in Arizona seems to represent a
relict population, most likely from a more extensive Ice Age distribution continuous with the Californian
populations. Similar Ice Age relict populations, on either side of the Salton Trough are commonplace (e.g..
Van Devender 1990, 2007; Felger & Van Devender 2010).

There are approximately 5 shrubby species of Salvia in Arizona: S. dorrii (Kellogg) Abrams complex
(includes S. pachyphylla Epling ex Munz), S. mohavensis Greene, S. parryi A. Gray, S. pinguifolia (Fernald)
Wooton & Standley, and S. vaseyi. Among these, S. vaseyi is the most xeric-inhabiting species and has the
narrowest ecological and geographic ranges and can be distinguished by its rugulose leaf blades with crenu-
late margins and bristle-tipped calyx teeth.

Baja California: summit of grade to Colorado Desert on road from Tia Juana to Mexicali. 14 May 1925, Peirson 5882 (RSA!); dry
slopes, 38 mi W of Mexicali, 2500 ft, 14 May 1925, Munz 9582 (RSA!); East slope of the Sierra Juarez along Hwy 2, steep slopes with

ACKNOWLEDGMENTS

The Study leading to this work was supported by the United States Fish and Wildlife Service, Arizona Game
and Fish Department, Foundation for North American Wild Sheep, Boone and Crockett Club, Desert Big-
horn Council, and Arizona Desert Bighorn Sheep Society. Felger thanks W. Eugene Hall and Philip David
Jenkins at University of Arizona; Andy Sanders at University of California, Riverside; Judy Ann Gibson at
San Diego; and Sula Vanderplank at RSA for generous assistance. Adrian Quijada-Mascarenas provided the

REFERENCES

Broyles, B., L. Evans, R.S. Felger and G.P. Nabhan. 2007. Our grand desert: a gazetteer for northwestern Sonora,
southwestern Arizona, and northeastern Baja California. In: R.S. Felger and B. Broyles, eds. Dry borders: great
Natural Reserves of the Sonoran Desert. University of Utah Press, Salt Lake City. Pp. 581-679.

Consortium of Caufornia Herbaria. 201 0. httpy/ucjeps.berkeley.edu/consortium/

Felger R.S., M. Wilson, K. Mauz, and S. Rutman. 2007. Botanical diversity of southwestern Arizona and northwestern

759

Journal of the Botanical Research Institute of Texas 4(2)

Sonora. In; R.S. Felger and B. Broyles, eds. Dry borders: great Natural Reserves of the Sonoran Desert. University
of Utah Press, Salt Lake City. Pp. 202-271 .

Felger, R.S. andT.R. Van Devendek in press. Flora ofTinajas Altas, Arizona— a century of botanical forays and forty
thousand years of Neotoma chronicles. Proc. San Diego Nat. Flist. Mus.

Hickman, J.C (ed.). 1 993. The Jepson manual. University of California Press, Berkeley.

Shreve, F. 1 951 .Vegetation of the Sonoran Desert. Carnegie Institution ofWashington Publication no. 591 . Wash-
ington, D.C. Reprinted as vol. I of Shreve & I.L. Wiggins, Vegetation and flora of the Sonoran Desert. Stanford
University Press, Stanford.

Southwest Environmental Information Network. 2009. Accessed 20 January 2010 from http/Zswbiodiversity.org/seinet/
index.php.

Turner, R.M. and D.E. Brown. 1 994. Sonoran Desertscrub. In: D.E. Brown, ed. Biotic communities of the southwestern
United States and northwestern Mexico. University of Utah Press, Salt Lake City. Pp. 1 81-221 .

Van Devender, T.R. 1990. Late Quaternary vegetation and climate of the Sonoran Desert, United States and Mexico.
In; J.L Betancourt, T.R. Van Devender, and PS. Martin, eds. Packrat middens: the last 40,000 years of biotic
change. University of Arizona Press, Tucson. Pp. 134-165.

Van DEVENDEFtTR. 2007. What packrats told us about deep ecology and the ecological detectives who solved the
case. In: R.S. Felger and B. Broyles, eds. Dry borders: great Natural Reserves of the Sonoran Desert. University
of Utah Press, Salt Lake City. Pp. 58-68.

Western Regional Cumate Center 201 0 www.wrcc.dri.edu/CLIMATEDATA.html

PASPALUM PUBIFLORUM AND P. QUADRIFARIUM (POACEAE)

NEW TO CALIFORNIA, WITH A KEY AND NOTES ON INVASIVE SPECIES

Richard E. Riefner, Jr.

Silvia S. Denham

Research Associate
Rancho Santa Ana Botanic Garden
1500 North College Avenue
Claremont, California 9171 1-3157, U.SA.
rriefner@earthlink.net

istituto de Botdnica Darwinian

J. Travis Columbus

1500 North College Avenue
California 91 71 1-31 57, 1
'is.coiumbus@cgu.edu

ABSTRACT

RESUMEN

INTRODUCTION

Paspalum L. (Poaceae, Panicoideae) is comprised of approximately 350 species, which are distributed pri-
marily in the subtropical, tropical, and warm-temperate regions of the Americas (Zuloaga & Morrone 2005;
Denham et al. 2010). However, owing to their utility as turf, forage or ornamental grasses, many taxa are
now widely distributed and have become some of the world’s most troublesome weeds (Holm et al. 1979;
Weber 2003; GCW 2010; PIER 2010).

The genus Paspalum, in general, is easily recognized by its unilateral racemes distributed along the
main axis of the inflorescence, plano-convex spikelets with the upper lemma oriented towards the rachis,
and the lower glume typically being absent. Confident, species-level determinations, however, often prove
difficult. Comparative ecological studies, such as growth responses to salinity and soil saturation, and veg-
etative morphology, phenology, and habitat associations can improve separation and help explain species

since weeds are generally under-collected and thus poorly represented in herbaria, the distribution of many
nonnative taxa in North America is poorly known (Allen & Hall 2003). Paspalum vaginatum Sw. (seashore
paspalum), native to the subtropical and tropical regions of the New World, has been introduced widely and
is now naturalized in warm, coastal regions around the world (Allen & Hall 2003; Weber 2003; Zuloaga et
al. 2003). Naturalized populations were reported for the first time for California by Riefner and Columbus
(2008).

762

In this paper, we provide the first documented records of P. pubijlorum Rupr. ex E. Foum. a
qmdrifarium Lam. for California; P. pubijlorum was collected from Orange and western Riverside cou
and P. quadrifarium from Los Angeles County in southern California. New records are also docum^
for P. vaginatum, which is invading estuarine and other saline wetlands in southern California. We n
the naturalized status, habitats occupied, mode of introduction, and provide a summary of the regie
California where recently introduced species of Paspalum have been observed or might become invas:
key is also provided to identify the known species of Paspalum in the State.

NEW RECORDS

CALIFORNIA

Paspalum pubijlorum (hairyseed paspalum) and P. quadrifarium (tussock paspalum) have not been reported
previously for California in treatments of the nonnative Poaceae growing outside of cultivation (Hitchcock
1951; Webster 1993; Bossard et al. 2000, 2006; Hrusa et al. 2002; Allen & Hall 2003; DiTomaso & Healy
2003, 2007; Roberts et al. 2004; Rebman & Simpson 2006; Bossard & Randall 2007; Clarke et al. 2007;
Grewell et al. 2007; Riefner & Boyd 2007; Dean et al. 2008; Roberts 2008; Jepson Flora Project 2010; USDA
2010a, b).

Paspalum pubijlorum is a perennial grass native to the eastern and southwestern United States (Pennsylvania
to Texas and Colorado), Mexico, and Cuba (Allen & Hall 2003; Denham et al. 2010; USDA 2010a). In the
United States, it grows on moist open ground and disturbed areas, in wet meadows, on banks and edges of
forests, streams, ponds, lakes, and irrigation ditches, especially in alkaline or calcareous soils (Hitchcock
1951; Correll & Correll 1975; Allen & Hall 2003).

Paspalum pubijlorum has branched decumbent culms that frequently root at the nodes (Fig. la). It has
(2-)3-12 racemes per inflorescence with pubescent, rarely glabrescent or glabrous paired spikelets, which
are elliptic to obovate-elliptic and (2.4-)2.6-3.2 mm long (Denham et al. 2010). The number of racemes
per inflorescence is not a fixed character and varies considerably; Hitchcock (1951) reports 3-5 racemes,
and Allen and Hall (2003) report 2-7 racemes per inflorescence. In southern California, populations typi-
cally have 3-6 racemes per inflorescence, but it is not uncommon to find plants having 2-3 racemes per
inflorescence (Fig. lb). Paspalum hartwegianum E. Foum. (Hartweg’s paspalum) is similar to P. pubijlorum. It
has simple erect culms with 3-23 racemes (also highly variable) per inflorescence, and paired pubescent
spikelets, which are elliptic to obovate and (2.3-)2.6-3.1 mm long (Denham et al. 2010). Accordingly, careful
collecting and documentation in the field is needed to accurately separate P. pubijlorum from P. hartwegianum
and other closely related species; currently, P. hartwegianum is not known to occur in California.

In southern California, P. pubijlorum could be confused with robust forms of P. distichum L. (knotgrass),
which is a native rhizomatous or stoloniferous perennial with spikelets pubescent only on the back of the
upper glumes. Its inflorescence is digitate (with two branches), but a third lower branch may occasionally
be present. Paspalum distichum is a highly variable species; it can be slender and creeping or robust and
cespitose. Although P. distichum usually has solitary spikelets, racemes with only paired spikelets or racemes
with paired and solitary spikelets can be present on the same plant. Robust forms of P. distichum with mostly
3 racemes per inflorescence and paired spikelets collected in southern California have been called Paspalum

paucispicatum Vasey (Hitchcock 1951). Paspalum paucispicatum (a synonym of P.

2003) has often been confused with P. pubiflorum (Verloove & Reynders 2007a).

Paspalum pubiflorum can be weedy when introduced to new regions (GCW 2009). In southern California,
it can form dense mats that compete for space with low-growing native hydrophytes in riverine and urban
creek habitats. Although P. pubiflorum occupies alkaline habitats, and unlike P. distichum and P. vaginatum,
which are known halophytes (Menzel & Lieth 2003), it is not expected to successfully colonize highly saline

in California is probably the result of ;

rge cespitose perennial native to Argentina, Brazil, Paraguay, and Uruguay (Al-
len & Hall 2003; Zuloaga et al. 2003). In Argentina, P. quadrifarium forms dense tussocks in the Flooding
Pampa grasslands, on river banks or lake shores, and is often regarded as a weed (Ortega & Laterra 2003;
Herrera et al. 2005).

eastern states (Garbari 1972; Allen & Hall 2003; USDA 2010b). It has been identified as a noxious weed or is a
potential invasive pest in New South Wales and Queensland, Australia, coastal Italy, and the southern United
States (Allen & Hall 2003; Verloove & Reynders 2007b; Bargeron et al. 2008; Sydney Weeds Committees
2010 ).

In southern California, P. quadrifarium most likely escaped from cultivated sources, and currently is known
only from a small population growing in disturbed riparian scrub. Accordingly, this species is here consid-
ered tenuously established and its dispersal and naturalization remains

University Dr., UTM (NAD 83) IIS
Rie/ner 09-10 (RSA). Riverside Co.:
D83)11S0550217E
t, 5Jul 2008. Rie/ner 08-215 (RSA). San Diego
(NAD 83) IIS 0464168E 3674334N, elev. ca.
08-288 (RSA).

Paspalum vaginatum is known from warm temperate, tropical, and subtropical regions around the world, and
is widely regarded as an invasive species; see Erickson and Puttock (2006), ISSG (2008), and Riefner and
Columbus (2008) for reviews. Worldwide, and outside of cultivation, P. vaginatum occupies coastal salt and
brackish water marshes, shallow-water lagoons and tidal channels, mangroves, coastal shrublands, dunes
and beaches, summer-moist saltpans, wet pastures, and freshwater riparian and floodplain habitats (Allen
& Hall 2003; Shaw & Allen 2003; Weber 2003; Siemens 2006). In southern California, rapidly expanding
populations and the formation of dense monocultures of P. vaginatum pose a serious threat to the structure,
function, and native species composition of estuarine wetlands (Riefner & Columbus 2008).

n control, and for turf, including specific
■ow & Duncan 1998; Duncan & Carrow

Riefner et al.. Two Paspalum species new for California

Paspalum vaginatum, a stoloniferous and rhizomai
bilitation of salt-affected lands, forage, dune stabilizatic
ecotypes and cultivars with improved tolerance to sali
1999; Duncan 2003). Paspalum vaginatum cultivars and ecotypes can maintain growth and vigor under ir-
rigation with seawater, i.e., seawater is approximately EC^ (electrical conductivity of water) 54 dS/m"‘ (deci-
Siemens per meter) or -34 ppt (parts per thousand) salt (Duncan & Carrow 1999; Lee et al. 2005; Berndt
2007; Pessarakli 2007). For comparison, freshwater habitats contain <0,5 ppt salt or <1 dS/m-‘ (Cowardin
et al. 1979); the minimum criterion required for a species to be classified as a halophyte is a salinity level
having an electrical conductivity measurement of at least 7-8 dS/m'^ during significant portions or all of
the plant’s life cycle (Aronson 1989).

Although we cannot identify specific P. vaginatum ecotypes/cultivars that have been introduced to
southern California or those growing outside of cultivation in wildlands, high-salinity tolerant plants are
now established in tidal wetland habitats, including sea beaches within the high tide zone (Fig. 2a). These

sibly establish as far north as central California.

In the coastal lowland wetlands of the Hawaiian Islands, P. vaginatum is highly invasive in brackish
wetlands (i.e., the mixohaline salinity classification [0.5-30 ppt salt] of Cowardin et al. 1979) (Bantilan-
Smith et al. 2009). Although P. vaginatum occupies a variety of wetland and saline environments in southern

vaginatum is highly invasive predominately in brackish wetland habitats.

As a result of continuing field documentation, unvegetated sand flats and mudflats located along tidal
creeks and lagoons at the head of coastal bays and estuaries appear most vulnerable to colonization by P.
vaginatum (Fig. 2b). Tidal mudflats are highly productive areas for invertebrates and provide rich foraging
habitat for shorebirds at low tide and other birds and fish at high tide (EPA 2010). Worldwide, invasions of P.
vaginatum are converting unvegetated or sparsely vegetated tidal flats and shallow lagoon habitats to vegetated

habitat and food resources^of shorebirds (Siemens 2006; ISSG 2008; Bird Life Imernational 2009).

Human activities such as dredging and filling have contributed to the loss of tidal mudflats in southern
California (Williams & Desmond 2001). Invasions by nonnative species, however, now also pose a threat
to estuarine wetland ecosystems (Grewell et al. 2007). Accordingly, development of urban watershed and
estuary conservation management plans should include identification and eradication programs to deter the
spread of nonnative plants, especially potentially invasive halophytes that could further degrade sensitive
estuarine wetlands in southern California.

s the Consortium of California Herbaria (2010), a

DISCUSSION

1 , and a review of pertinent literature and electronic database:
summary of the distributional records
current naturalized status, occupied habitats, document invasive behavior, and speculate where recentl)
introduced Paspalum species might become invasive. These data are summarized in Table 1. Naturalizatior
categories shown in Table 1 are somewhat subjective, but follow Hrusa et al. (2002) in order to allow con-
sistency in data compilation of new

Dean et al. (2008) recently pointed out the confusion regarding identifications and the relal
ized distributions of Paspalum notatut

Considering the ongoing difficulties with separating P. distichum from P. vaginati
key that will serve to identify the species and varieties of Paspalw

Parodi in California,
provide the following
occur in California.

Journal of the Botanical Research Institute of Texas 4(2)

ACKNOWLEDGMENTS

We greatly appreciate the efforts of Harvey Brenneise and Irene Holiman (Library of Rancho Santa Ana

Botanic Garden) for assistance with document retrieval, and to Garn Wallace (Wallace Laboratories, El

Segundo, California) for helpful discussion of salinity analysis and classifications. Fred Hrusa, California

Department of Food and Agriculture (CDA), and an anonymous reviewer provided helpful comments that

from the International Foundation for Applied Research in the Natural Sciences (IFARNS) to study alkaline al-
luvial habitats in coastal southern California.

REFERENCES

Allen, C.M. and D.W. Hall. 2003. Paspatum. In: Flora of North America Editorial Committee, eds. Flora of North
America north of Mexico, vol. 25, Magnoliophyta: Commelinidae (in part): Poaceae, part 2. Oxford University
Press, New York, NY. Pp. 566-599.

Aronson, J.A. 1 989. Haloph: a data base of salt tolerant plants of the world. Office of Arid Land Studies, University
of Arizona, Tucson.

Bargeron, C.T., C.R. Minteer, C.W. Evans, DJ. Moorhead, G.K. Douce, and R.C. Reardon (technical coordinators). 2008.
Invasive plants of the United States: identification, biology and control (DVD-ROM). USDA Forest Service
Technology Enterprise Team, FHTET-08-1 L Morgantown, WV.

BANTiLAN-SMrTH, M., G.L. Bruland, R.A. Mackenzie, A.R. Henry, and C.R. Ryder 2009. A comparison of the vegetation and
soils of natural, restored, and created coastal lowland wetlands in Hawaii. Wetlands 29:1023-1035.

Berndt, W.L Salinity affects quality parameters of 'SeaDwarf seashore paspalum. HortScience 42:41 7-420.

Bird Life International 2009. Important bird area factsheet: Rietviei Wetland Reserve, South Africa. Available:
httpY/www.birdlife.org/datazone/sites/index.html?action=SitHTMDetails.asp&sid=7165&m=0 [accessed
January 2010].

Bossapd, C M. Brooks^ JM DTomaso, JM. Rancwi, C Fk>L 1 Sioq A Stanicn, and R Warner 2006. California invasive plant inven-
tory. California Invasive Plant Council, Publ. No. 2006-02, Berkeley.

Bossard, C.C., J.M. Randall, and M.C Hoshovesky. 2000. Invasive plants of California's wildlands. University of Cali-
fornia Press, Berkeley.

Carrow, R.N. and R.R. Duncan. 1998. Salt-affected turfgrass sites: assessment and management. Ann Arbor Press,
Chelsea, ML

Clarke, O.F., D. Svehla, G. Ballmer, and A. Montalvo. 2007. Flora of the Santa Ana River and environs. Heyday Books,
Berkeley.

Consortium of California Herbaria. 2010. Paspalum. Available: httpY/ucjeps.berkeley.edu/consortlum/ [accessed
May-June 2010].

Correu, D.S. AND H.B. CoRRELL 1975. Aquatic and wetland plants of southwestern United States, vol.1. Stanford Uni-
versity Press, Stanford, CA.

CowARDfrj, LM., V. CARTEft F.C GoLET, AND E.T. LaRoe. 1 979. Classification of wetlands and deepwater habitats of the United
States. U.S. Department of the Interior, Fish and Wildlife Service, Office of Biological Services, FWS/OBC-79/31,
Washington, D.C.

Dean, E., F. Hrusa, G. Leppig, A. Sanders, and B. Ertter 2008. Catalogue of nonnative vascular plants occurring spon-
taneously in California beyond those addressed in The Jepson Manual-Part II. Madrono 55:93-1 1 2.

Denham, 5.S., O. Morrone, and F.O. Zuloaga. 201 0. Estudios en el genero Paspalum (Poaceae, Panicoideae, Paniceae):
Paspalum denticulatum y especies afines. Annals of the Missouri Botanical Garden 97:1 1-33.

DTomaso, J.M. and EA. Healy. 2003. Aquatic and riparian weeds of the West. U.C Agriculture and Natural Resources
Publication 3421, Oakland, CA.

DTomaso, JJvl. AND EA. Healy. 2007. Weeds of California and other western states, vol. 2, Geraniaceae-Zygophyllaceae.
U.C. Agriculture and Natural Resources Publication 3488, Oakland, CA

Duncan, R.R. and RN. Carrow. 1 999. Seashore paspalum:the environmental turfgrass. John Wiley & Sons, Inc, Hoboken, NJ.

770

Siemens, TJ. 2006. Impacts of the invasive grass saltwater paspalum {Paspalum vaginatum) on aquatic com-
munities of coastal wetlands on the Galapagos Islands, Ecuador. Master of Science Thesis, Cornell University,
Ithaca, NY.

Sydney Weeds Committees. 2010. Noxious weeds. Available: httpy/www.sydneyweeds.org.au/noxious-weeds.php
[accessed January 2010].

United States Department of Agriculture (USDA). 201 Oa. PLANTS database: plants profile for Paspalum pubiflorum Rupr.
ex E. Fourn., hairyseed paspalum. United States Department of Agriculture, Natural Resources Conservation
District. Available: httpY/plants.usda.gov/java/profile?symbol=PAPU5 [accessed January 2010].

United States Department of Agriculture (USDA). 201 Ob. PLANTS database: plants profile for Paspalum quadrifarium
Lam., tussock paspalum. United States Department of Agriculture, Natural Resources Conservation District.
Available: httpy/plants.usda.gov/java/profile?symbol=PAQU6 [accessed January 2010].

United States Environmental Protection Agency (EPA). 2010. Wetlands of the Pacific Southwest: mudflats. Available:
httpY/www.epa.gov/region09/water/wetlands/mudflats.html [accessed January 2010].

Verloove, F. and M. Reynders. 2007a. Studies in the genus Paspalum (Paniceae, Poaceae) in Europe-1 . Pospa/um
distichum subsp. paucispicatum, an overlooked taxon in France. Willdenowia 37:199-204.

Verloove, F. and M. Reynders. 2007b. Studies in the genus Paspalum (Paniceae, Poaceae) in Europe-2, the Quadrifaria
group. Willdenowia 37:423-430.

Weber, E. 2003. Invasive plants of the World. CABI Publishing, CAB International, Wallingford, UK.

Webster, R. 1993. Paspalum. In: J.C. Hickman, ed.The Jepson manual: higher plants of California. University of
California Press, Berkeley. P. 1 280.

Williams, G.D. and J.S. Desmond. 2001 . Restoring assemblages of invertebrates and fishes. In: J.B. Zedler, ed. Handbook
for restoring tidal wetlands. CRC Press, Boca Raton, London, New York, Washington, D.C Pp. 235-269.

ZuLOAGA, F.O. and O. Morrone. 2005. Revision de las especies de Paspalum para America del Sur austral (Argentina,
Bolivia, sur Del Brasil, Chile, Paraguay y Uruguay). Monogr. Syst. Bot. Missouri Bot. Card. 1 02:1 -297.

ZuLOAGA, F.O., O. Morrone, G. Davidse,T.S. Filgueiras, P.M. Peterson, RJ. Soreng, and E.J. JuoziEwia. 2003. Catalogue of
New World grasses (Poaceae): III. Subfamilies Panicoideae, Aristidoideae, Arundinoideae, and Danthonioideae.
Contr.US. Natl. Herb. 46: 1-662.

SARRACENIA MINOR VAR. OKEFENOKEENSIS (SARRACENIACEAE)
DISCOVERED OUTSIDE OF THE OKEFENOKEE SWAMP AREA

Jacob S. Thompson

Georgia Department of Natural Resources
Wildlife Resources Division, Nongame Conservation Seaion
Brunswick, Georgia 3 1520, U.S.A
jacob.thompson@dnr.state.ga.us

Sarracenia minor Walt. var. okefenokeensis Schnell (Okefenokee Giant) was first recognized as a new variety
of Sarracenia minor in 2002 and is currently defined as endemic to the Okefenokee Swamp area in southeast
Georgia (Schnell 2002; Weakley 2010). The Okefenokee variety differs from S. minor var. minor in morphol-
ogy, habitat requirements, and flowering time (Schnell 2002; NatureServe 2010). Individuals of S. minor var.
okefenokeensis are generally much taller than S. minor var. minor, averaging 70-90 cm in height (vs. 25-35
cm in var. minor) and have a more slender appearance. Furthermore, S. minor var. okefenokeensis flower about
two weeks later at the same latitude and prefer a much wetter habitat. These differences are maintained in
a common garden (Schnell 2002).

Sarracenia minor var. okefenokeensis has been designated by NatureServe as G4T2T3 (Globally Imperiled)
and has a state rarity rank of S2S3 (Imperiled) in Georgia (NatureServe 2010). Previous work has suggested
that populations may be found from 5 km (Schnell 2002) to 8 km (NatureServe 2010) outside the borders
of the Okefenokee National Wildlife Refuge (NWR). Unlike the protected populations that exist within
the borders of Okefenokee NWR, populations in surrounding areas may suffer from a variety of different
anthropogenic threats (NatureServe 2010). Also, there is a lack of population locality information.

The novel occurrence reported here was found 30 km northeast of the Okefenokee Swamp border
across the Satilla River (Fig. 1), though in a basin swamp environment similar to those within the Okefeno-
kee. A population of S. minor var. okefenokeensis was found in Kings Bay, a 1390 ha peat-filled nonriverine
basin swamp that stretches 13 km from southeast Brantley County to northwest Camden County (not to be
confused with Kings Bay Naval Base in Camden County). Kings Bay exists on a Pleistocene barrier island
and was probably a large marshy tidal lagoon in ancient times. Currently, Kings Bay and the Okefenokee
Swamp are not connected and it is uncertain whether they were joined in previous times. Thousands of
S. minor var. okefenokeensis individuals were found on floating Peatmoss (Sphagnum spp.)
wardia virginica, Peltandra sagittifolia, Nymphaea odorata ssp. odorata, Lachnanthes caroliniam
Rhynchospora distans, and Utricularia subulata. The Peatmoss mats were floating o
of water. Plants were also found on scattered hummocks with shrubs of Lyonia lucida <
laurifolia. Small trees of Pinus elliottii var. elliottii, Taxodium ascendens, and Gordonia las
spaced throughout the community.

Voucher specimen: GEORGIA. Brantley Co.: Kings Bay, S of Kings Bay Rd.. 4 km NW of the Kings Bay Rd ai

J. Bot Res. Inst Texas 4(2): 771-77

774

BOOK REVIEW

Jim Stanley 2009. Hill Country Landowner’s Guide. Louise Lindsey Merrick natural environment series,
no. 44. (ISBN 978-1-603-44137-7, flexbound w/flaps). Texas A &r M Press, John H. Lindsey Building,
Lewis Street, 4354 TAMU, College Station, Texas 77843-4354, U.S.A. (Orders; www.tamupress.com,
1-800-826-8911). $19.95, 224 pp., 40 color photos, 1 map, 5 x 8 W.

POLYCARPON TETRAPHYLLUM (CARYOPHYLLACEAE)
NEW TO THE FLORA OF LOUISIANA

Charles M. Allen, Jarrod Grandon, Krisztian Megyeri, and Brad Waguespack

Colorado State University
Fort Polk Station 1645 23rd St
Fort Polk, Louisiana 71459, U.S.A

member of the Caryophyllaceae. It is characterized by mostly whorled leaves, keeled sepals 1.5-2. 5 mm
long, and stipules 1.8-2. 8 mm long (Thieret & Rabeler 2005). The other species of Polycarpon in the US is
California manyseed (P. depressum Nutt.) with opposite not whorled leaves, flat sepals 1-1.5 mm long, and
stipules 0.4-1. 2 mm long. Four-leaved manyseed is a native of the Mediterranean region of southern Europe
and has been introduced into Ala., Calif., Fla., Ga.. Ore., S.C., and Tex. in the United States and also British
Columbia (Thieret & Rabeler 2005; USDA NRCS 2010). There are also historic records from Massachusetts
and Pennsylvania. It is not listed for Louisiana by Thomas and Allen (1996) nor in the Plants database (USDA
NRCS 2010) so our collection is apparently the first for the state.

REFERENCES

Thieret, J.W. and R.K. Rabeler. 2005. Polycarpon. In: Flora of North America Committee, eds. Flora of North America
north of Mexico. Vol. 5. Oxford University Press, New York. Pp. 25-26.

Thomas, R.D. and CM. Allen. 1996. Atlas of the vascular flora of Louisiana, Vol. 2: Dicotyledons Acanthaceae-
Euphorbiaceae. Louisiana Department of Wildlife and Fisheries, Baton Rouge, LA.

USDA, NRCS. 2010. The PLANTS database (http-y/plants.usda.gov/plants). National Plant Data Center, Baton
Rouge, LA 70874-4490.

776

BOOKS RECEIVED

Lane Greer and John M. Dole. 2009. Woody Cut Stems for Growers and Florists: How to Produce and
Use Branches for Flowers, Fruit and Foliage. (ISBN 978-0-81-92892-1, hbk.). Timber Press, Tbe
Hastings Building, 133 S.W Second Avenue, Suite 430, Portland, Oregon 97204-3527, U.S.A. (Orders:
orders@timeberpress.com, 1-800-827-5622). $39.95, 576 pp., 137 color pbotos and nine line draw-
ings, 6" X 9".

Patricia A. Harding. 2008. Huntleyas and Related Orchids. (ISBN 978-0-881-92884-6, bbk.). Timber
Press, Tbe Hastings Building, 133 S.W. Second Avenue, Suite 450, Portland, Oregon 97204-3527,
U.S.A. (Orders: orders@timeberpress.com, 1-800-827-5622). 39.95, 260 pp., 150 color pbotos and
five line drawings, 7y8" x lOYs".

Mary S. G. Lincoln 2008 Uverworts of New England: A Guide for the Amateur Naturalist. (ISBN 978-
0-893-27478-8, bbk.). Memoirs of tbe New York Botanical Garden, Vol. 99. Tbe New York Botanical
Garden Press, 200tb Street and Kazimiroff Boulevard, Bronx, New York 10458-5126, U.S.A. (Orders:
bttp://nybgpress.com, nybgpress@nybg.org, 1-718-817-8721, 1-718-817-8842 fax). $45. 00; 162 pp.;
color pbotos, line drawings, and distribution maps; 7V4" x 10y4".

James W Hinds AND Patricia L. Hinds. 2007. MacroUchens of New England. (ISBN 978-0-893-27477-1, bbk.).
Memoirs of tbe New York Botanical Garden, Vol. 96. Tbe New York Botanical Garden Press, 200tb Street
and Kazimiroff Boulevard, Bronx, New York 10458-5126, U.S.A. (Orders: bttp://nybgpress.com, nybg-
press@nybg.org, 1-718-817-8721, 1-718-817-8842 fax). $65.00, 608 pp., color pbotos, 7%" x IO1/4".

Charies R. Hatch. 2007. Trees of the California Landscape. (ISBN 978-0-520-25 124-3, bbk.). University of
California Press, Berkeley, California 94704, U.S.A. (Orders: www.ucpress.edu or Califomia-Princeton
Fulfillment Services, 1445 Lower Ferry Road, Ewing, New Jersey 08618, U.S.A., 1-609-883-1759, 1-609-
883-7413 fax). $60.00, 1000+ illustrations: color pbotos, drawings, and maps, 8%" x iP/s".

Tim Robinson 2008. William Roxburgh: The Founding Father of Indian Botany. (ISBN 978-1-860-
77434-2, bbk.). Pbillimore & Co. Ltd., Dene Road, Healey House, Andover, Hampshire, SPIO 2AA,
England, U.K. (Orders: www.pbillimore.co.uk or publisbing@pbillimore.co.uk, 44-01264-409200).
£50.00, 286 pp.,8W’xll".

Matthew A. JENKS AND Andrew J. Wood (eds). 2007. Plant Desiccation Tolerance. (ISBN 978-0-813-81263-2,
bbk.). Wiley, 10475 Crosspoint Boulevard, Indianapolis, Indiana 46246, U.S.A. (Orders: www.wiley.
com, 1-800-762-2974, 1-800-597-3299 fax). $209.99, 311 pp., 7" x lOW'.

Peter Mickulas. 2007. Britton’s Botanical Empire: The New York Botanical Garden and American
Botany, 1888-1929. (ISBN 978-0-893-27479-5, bbk.). Tbe New York Botanical Garden Press, 200*
Street &r Kazimiroff Boulevard, Bronx, New York 10458-5126, U.S.A. (Orders: www.nybgsbop.org or
customerservice@nybgsbop.org, 1-718-817-8869). $45.00, 328 pp., 7" x 10.5".

Andrew Brown, Pat Dundas, Kingsley Dixon, and Stephen Hopper. 2008. Orchids of Western Australia. (ISBN
978-0-980-29645-7, bbk.). University of Western Australia Press, 35 Stirling Highway, Crawley, Perth,
Western Australia 6009. (Orders: www.uwap.edu.au or admin@uwap.uwa.edu.au, 61-8-6488-3670,
Aus. $89.95, 421 pp., 9%" x UVz”.

61-8-6488-1027 fax), j

ADDITIONS TO THE VASCULAR FLORA OF NEW MEXICO

Ben S. Legler

WTU Herbarium
University of Washington
Box 355325

Seattle, Washington 98195-5325, USA
blegler@u.washington.edu

ABSTRACT

States (Stein 2002), with Allred (2009) reporting 3,238 species. However, the cataloging of the state’s flora

state records awaited discovery in New Mexico. This has been confirmed by the 417 state records reported
in the New Mexico Botanist Newsletters (http://aces.nmsu.edu/academics/rangescienceherbarium/the-new-
mexico-botanist-.html) from January, 1996 to February, 2010. Documented here are 25 more additions to
the state’s flora. Most of these were uncovered during a floristic inventory of Vermejo Park Ranch (VPR) from
2007 to 2009 in Colfax and Taos Counties (Legler 2010). The remainder were found during field surveys in
New Mexico for Botrychium in 2009. All specimens are deposited at the Rocky Mountain Herbarium (RM)
with duplicates, where available, at the University of New Mexico Herbarium (UNM). Additional duplicates

Allred (2009), and secondarily on BONAP (2010), the USDA Plants Database (NRCS 2010), Fima of North
America (FNA 1993+), and regional floras. Identifications for all cited specimens were verified by B.E. Nelson
(Herbarium Manager, RM) or by specialists as listed below. Two taxa not included here, Chionophila jamesii
Benth. and Platanthera obtusata (Banks ex Pursh) Lindl. ssp. obtusata, were recently reported as new for the
state by Peterson (2010) and Heil (2009), respectively. Additional collections of these were obtained during
the inventory of VPR. Their label data are available through the RM web site (Hartman et al. 2009).

ASTERACEAE

Heterotheca pumila (Greene) Semple — This endemic of subalpine and alpine habitats in Colorado, Utah, and
Wyoming has been erroneously reported for New Mexico (e.g., BONAP 2010). According to Semple (2006),
“Reports of occurrence of H. pumila from Arizona and New Mexico are based on narrow-leaved plants of H.
fulcrata var. amplifolia with small ovate-lanceolate bracts subtending the heads.” The collection cited here is

J. Hot Res. Inst. Teas M2): 777 -m. 2010

fulcrata var. amplifolia. Its morphology matches H. pumila, with the leaves oblanceolate throughout

e upper leaves immediately subtending and greatly surpassing the solitary heads.

specimen: Taos Co.: VPR: 1.2 air mi S of Big Costilla Peak and 0.1 air mi E of point 12931, N36.90328, W105.33025, 12,542

BRASSICACEAE

Draba streptobrachia R. A. Price — ^This species was considered endemic to the high mount
(Al-Shehbaz et al. 2010). Verified by Ihsan Al-Shehbaz, Missouri Botanical Garden (MO).

CYPERACEAE

Carex microglochin Wahlenb. ssp. microglochin— Cochrane (2003) states that this species is “usually

documented as far south as Colorado (BONAP 2010). It is a highly distinctive but easily overlooked sedge of
bogs, fens, and other peaty or wet habitats. Verified by Peter Zika, University of Washington (WTU).

Carex nelsonii Mack. — ^This Rocky Mountain endemic ranges from southern Montana south to Colorado
and Utah (Murray 2003). Although not previously reported from New Mexico, it has been documented from
several bordering counties in Colorado (BONAP 2010). Verified by Zika.

Voucher specimen: Taos Co.: VPR: 0.95 air mi S of Big Costilla Peak, N36.90656, W105.32567, 12,234 ft elev., flowing snowmelt spring
on gravelly alpine flat, 25 Jul 2007, Legler 6514 (RM, UNM).

arctic-alpine species as extending

n fen, 29 Jul 2007, Legler 6665 (RM, I

n partly shaded by Picea, 19 Aug 2008, Legler 10939 (RM, UNM).

Jnncus biglumis L.— The cited collection represents a southern range extension of about 180 miles (290
km) from the nearest populations in Summit County, Colorado (BONAP 2010) for this circumpolar, arctic
species. The plants were rare and local at this site.

Juncus parryi Engelm. — ^The previously known distribution of this species included all states in or wesi
of the Rockies except for Arizona and New Mexico (Brooks & Clements 1993; BONAP 2010). Although ii

779

was not previously reported for New Mexico, one small, immature specimen at UNM appears to be this
species (Santa Fe Co.: Sangre de Cristo Mountains, Puerto Nambe Ridge below Pecos Baldy along Wind-
sor Trail, T18N RllE S2 NWV4, 10,800 ft elev., 17 Jul 1997, R. C. Sivinsfei 3920). All other specimens were
verified by Zika.

e, 25 Jul 2007, Legler 6548 (RM); VPR: slope oi

i SSW of Big Costilla Peak. N36.91531, W105.33058, 12,605 ft elev., rock outcrop
Wside of Lake No. 2. 1.3 air mi SSW of State Line Peak, N36.97764, W105.30591,
with scree and talus, 17 Jul 2008, Legler 9835 (RM. UNM).

Juncus triglnmis L. var. triglumis— Both varieties of this primarily arctic and boreal species occur in
North America. They extend south through the Rockies in scattered locations to Colorado for var. triglumis
and New Mexico for var. albescens Lange Fernald (Brooks & Clements 1993; BONAP 2010). An examina-

tion of specimens at RM showed many intermediat
differentiated. The most reliable characters appear t
perianth and the shape of the mature capsule apex. i
triglumis (.Legler 6699 and 10932) and one intermedial
J0986). Verified by Zika.

) be the length of the mature capsules relative to the
ited here are two collections clearly assignable to var.
: specimen that appears closest to var. triglumis (Legler

.1 airmiNE of Little Costilla Peak. N36.83944, W105.20361, 10,840

1, 29 Jul 2007, Legler 6699 (RM, 1

UNM); same site as previous, 19 Aug 2008, Legler 10932
of Big Costilla Peak, N36.95589, W105.31546, 11,962 ft

780

Endangered Species Act (USFWS 2001). It is widely scattered throughout boreal and western North America,
usually in very small populations (Farrar 2005), but is now known to be more common than previously
thought in Colorado (S. Popovich, pers. comm.). The plants cited here lit the unpublished B. Jurcatum',
currently considered indistinct from B. lineare (Farrar, pers. comm.). Forty-eight plants were counted at this
site. Verified by Farrar using enzyme electrophoresis.

Botrychium minganense Viet.— This widely distributed moonwort extends south throughout the western
cordillera to California, Arizona, and Colorado (Farrar 2005). As with B. hesperium, it is among the more
common moonwort species in adjacent Colorado (Popovich, pers. comm.) and has been overlooked in New
Mexico. Forty-eight collections of this species were obtained in New Mexico. Thirteen of these are cited here
to illustrate its distribution in the state. Data for the remaining vouchers are available through the RM web
site (Hartman et al. 2009). Populations at several sites numbered in the hundreds of plants. All collections
were verified by Farrar, including several by enzyme electrophoresis.

ORCHIDACEAE

Listera borealis Morong — ^The collection
cies south by about 160 miles (260 km) ft
Colorado (CONHP 2010). Plants were loct

air mi ENE of Costilla Reservoir dam. N36.88364,
tossy soil, 30 Jun 2007, Legler 5324 (RM); Carson
i6 crosses river, N36.56988, W105.38226, 10,611
?, Legler 11558 (RM).

Festuca hallii (Vasey) Piper — The large spikelets of this grass, with the glumes equaling or slightly e
ing the lemmas, are distinctive. The species is primarily distributed in the northern Great Plains of C
and the United States (Darbyshire & Pavlick 2007) but extends south in Colorado nearly to the New ^
border (Snow 2008; Hartman et al. 2009; BONAP 2010).

Ptilagrostis porteri (Rydb.) W.A. Weber — Previously thought to be endemic to central Colorado, this spe-
cies was considered for listing under the Endangered Species Act (USFWS 2005). The collection cited here
represents a southern range extension of about 150 miles (240 km) from the nearest populations in El Paso
County. Only a few dense clumps were observed at the location cited here. The plants were confined to the
tops of mossy hummocks in a habitat similar to known occurrences in Colorado.

Voucher specimen: Colfax Co.: VPR: near N end of Elk Meadows, 1. 1 air mi NE of Little Costilla Peak. N36.83899, W105. 20400, 10,880
ft elev., hummocky fen partly shaded by Picea, 19 Aug 2008, Legler 10940 (RM. UNM).

782

Journal of the Botanical Research Institute of Texas 4(2)

POLEMONIACEAE

Polemonium occidentale Greene var. occidentale — ^This widespread species has been documented from all
states in and west of the Rockies except for New Mexico (BONAP 2010). While examining specimens at UNM,
two collections of this species were found from Rio Arriba County (cited below) that had been misidenti-
fied. Additional collections from recent floristic inventories, also cited below, along with observations in the
field, suggest the species is not uncommon in wet mountain meadows in the northern portion of the state.

ocks,29Jul2007,Legler

6697 (RM, UNM). Rio Arriba C
2.1 air mi ENE of Costilla Reser

POLYGONACEAE

Eriogonum arcuatum Greene var. xanthum (Small) Reveal— This showy, mat-forming buckwheat was
previously considered endemic to the mountains of Colorado (Reveal 2005).

Voucher specimens; Taos Co.: VPR: in talus bowl 1.8 air mi S of Big Costilla Peak. N36.89408. W105.33172, 12,129 ft elev. northea.st-
facing, rocky, alpine slope. 19 Jul 2007, Legler 6310 (RM. UNM); VPR: 1.2 air mi S of Big Costilla Peak and 0.1 air mi E of point 12,931,
N36.90328, W105.33025, 12,542 ft elev., steep talus, scree, and rock slopes, 21 Jul 2007, Legler 6367 (RM); VPR: 0.23 air mi NE of Big
Costilla Peak. N36.92347, W105.32619, 12,172 ft elev., rocky, alpine slope. 18 jun 2008, Legler 8882 (RM. UNM); VPR: between 3rd and
4th lakes at Casias Lakes, 1.6 air mi S of Big Costilla Peak, N36.89698, W105.32964, 11,900 ft elev., dry, gravelly, subalpine meadow,
30 Jul 2008, Legler 10094 (RM).

. Three collections

_ i (1950) included

tion of this widespi

Phipps (1998), Phipps and O’Kennon (2004), and Allred (2009),

L Taber Morey Canyon 0.8 air mi W of Jones Canyon and 12 air mi WNW of

i, 2 Jun 2008, Legler 8350 (RM, UNM); VI

'NW of Raton. N36.96043, W104.66972, 7,767 ft elev., dry canyon bottom. 3 Jun 2008, Le^er
air mi SE of Long Canyon and 14.3 air mi WNW of Raton, N36.97458, W104.68166, 7,354 ft
tream, 3 Jun 2008, Ugler8387 (RM, UNM).

Potentilla nivea L.— The distribution of this circumpolar, arctic-alpine species extends south through the
Rockies to Utah and Colorado (Hulten 1968; BONAP 2010). It had been reported for New Mexico only by
Welsh (1982) and Welsh et al. (2003), but in both cases without documentation. Peterson (2000) questioned
Welsh’s report, and the species was not attributed to the state by Allred (2009). The collection cited here

Legler, Additions to the New Mexico flora

ACKNOWLEDGMENTS

The author thanks Vermejo Park Ranch, the New Mexico Native Plant Society, and the University of Wyoming
for supporting field work. Herbarium research was conducted at RM, UNM, and the University of Northern
Colorado Herbarium (GREE). The following individuals provided assistance and comments: Kelly Allred,
Donald Farrar, Ronald L. Hartman, Ken Heil, B.E. Nelson, Steve Popovich, Bob Sivinski, and Neil Snow.
I also thank Jill Larson for allowing me to cite a specimen of Polemonium occidentale collected during her
floristic inventory of the Carson National Forest.

REFERENCES

Al-Shehbaz, la., M.D. Windham, and R. Elven. 2010. Draba. In: Flora of North America Editorial Committee, eds. Flora
of North America north of Mexico. Vol. 7. Oxford University Press, New York, New York.

Allred, K.W. 2009. Flora Neomexicana I: the vascular plants of New Mexico. Published by the author (available
at httpy/www.lu lu.com/).

Ball P.W. 2003. Kobresia. In: Flora of North America Editorial Committee, eds. Flora of North America north of
Mexico. Vol. 23. Oxford University Press, New York, New York.

Ball, P.W. and D.E. Wujek. 2003. Eriophorum. In: Flora of North America Editorial Committee, eds. Flora of North
America north of Mexico. Vol. 23. Oxford University Press, New York, New York.

Biota of North America PraDCRAM (BONAP) 2010. U.S. county-level atlas of the vascularflora of North America. Floristic
Synthesis of North America. httpV/www.bonap.org/MapSwitchboard.html (accessed 4 Mar 2010).

Brooks, R.E. and S.E. Clemants. 1993. Juncus. In: Flora of North America Editorial Committee, eds. Flora of North
America north of Mexico. Vol. 22. Oxford University Press, New York, New York.

Cochrane, T.S. 2003. Carex sect. Leucoglochin. In: Flora of North America Editorial Committee, eds. Flora of North
America north of Mexico. Vol. 23. Oxford University Press, New York, New York.

Colorado Natural Heritage Program (CONHP). 201 0. Conservation status handbook (tracking lists). httpV/www.
cnhp.colostate.edu/download/list.asp (accessed 31 Jan 2010). Last updated 7 Jan 2010.

Darbyshire, SJ. and L.E. Pavlick. 2007. Festuca. In: Flora of North America Editorial Committee, eds. Flora of North
America north of Mexico. Vol. 24. Oxford University Press, New York, New York.

Farrai^ D. 2005. Moonwort systematics. Ayda Hayden Herbarium, Iowa State University, httpy/www.publiciastate.
edu/~herbarium/botrychium.html (accessed 16 Nov 2007).

Fernald, M.L 1950. Gray's manual of botany, 8th ed. D.Van Nostrand Co., New York, New York.

Flora of North America Editorial Committee, ed. (FNA). 1 993+. Flora of North America north of Mexico. Oxford Uni-
versity Press, New York, New York.

Hartman, R.L., B.E. Nelson, and B.S. Legler. 2009. Rocky Mountain Herbarium plant specimen database. httpv7www.
rmh.uwyo.edu/.

Heil K.D. 2009. Platanthera obtusata (Plant Distribution Report). In: New Mexico Botanist Newslett. Vol. 46. New
Mexico State University, Las Cruces, NM. httpy/aces.nmsu.edu/academics/rangescienceherbarium/the-new-
mexico-botanist-.html (accessed 10 Mar 2010).

HultEn, E. 1968. Flora of Alaska and neighboring territories. Stanford University Press, Stanford, California.

LEGLEit B.S. 2010. A floristic inventory of Vermejo Park Ranch, New Mexico and Colorado, M.S. Thesis, University
of Wyoming, Laramie, WY.

Martin, W.C. and RA COLEMAN. 2003. Listera. In: Flora of North America Editorial Committee, eds. Flora of North
America north of Mexico. Vol. 26. Oxford University Press, New York, New York.

Martin, W.C and C.R. Hutchins. 1 981 . A flora of New Mexico. 2 vols. Koenigstein, Germany.

Murray, D.F. 2003. Carex sect. Racemosae. In: Flora of North America Editorial Committee, eds. Flora of North
America north of Mexico. Vol. 23. Oxford University Press, New York, New York.

BOOK REVIEW

Journal of the Botanical Research Institute of Texas 4(2)

ANNOUNCEMENTS

2010 Delzie Demaree Travel Award Recipients
The 22nd Annual Delzie Demaree Travel Award was presented at the 57th Annual Systematics Symposium
(15-16 Oct 2010) at the Missouri Botanical Garden, St. Louis. Three students were presented the Travel
Award; Aliya Donnell, Ohio University, Laia Barres, The Morton Arboretum, and Alison Scott, University
ofWisconsin-Madison.

The 2010 Travel Awards were underwritten by 1) Delzie Demaree Travel Award Endowment, 2) Mem-
bers of the Delzie Demaree Travel Award Committee, and 3) John Clayton Chapter of the Virginia Native
Plant Society.

Anyone interested in making a contribution to Delzie Demaree Endowment Fund, which supports the
travel award, may make contributions by VISA or MasterCard or by a check, payable to Botanical Research
Institute of Texas, to Barney Lipscomb, 500 E 4th Street, Fort Worth, TX 76102-4025, U.S.A. 1-817-332-
7432; Email; barney@brit.org. Thank you.

The 201 1 Applications for the Delzie Demaree Travel Award
Applications for the 2011 Delzie Demaree Travel Award should include a letter from the applicant telling
how symposium attendance will benefit his/her graduate work and letter of recommendation sent by the
major professor. Please send letters of application to; Dr. Donna M.E. Ware, Herbarium, Biology Depart-
ment, The College of William and Mary, Williamsburg, VA 23187-8795; email: dmeware@verison.net. The
period for receiving applications will end three weeks prior to the date of the symposium if a sufficient
number of applications are in hand at that time. Anyone wishing to apply after that date should inquire
whether applications are still being accepted before applying. The Systematics Symposium dates for 2011
are 7-8 October 2011.

The Delzie Demaree Travel Award was established in 1988 honoring Delzie Demaree who attended
35 out of a possible 36 symposia before he died in 1987. Delzie Demaree was a frontier botanist, explorer,
discoverer, and teacher. His teaching career as a botanist began in Arkansas at Hendrix College in 1922. He
also taught botany at the University of Arkansas, Navajo Indian School, Yale School of Forestry, Arkansas
A&M, and Arkansas State University at Jonesboro where he retired as professor emeritus in 1953. One of
the things he enjoyed most as a botanist was assisting students with their field botany research.

( 2010 )

788

I new connbination in Lagotis (Plantaginaceae) by
David F, Murray, Reioar Elven, and Kanchi N. Gandhi

— 4(1):219

^ new combination in Lolium perenne (Poaceae:

A new Leucophyllum (Scrophulariaceae) from
Sonora, Mexico by James Henrickson andThomas R. Van
Devender— 4(2):581

A new Lomatium (Apiaceae) from the Ochoco
Mountains of central Oregon by Richard Helli-
WELL— 4(1):7

A new species of Angelonia (Plantaginaceae) from
Mexico by Kerry Barringer— 4(1 ):51

A new species of Calceolaria (Calceolariaceae) from
disturbed paramos in south Ecuador by Pamela
Puppo— 4{1):33

A new species of Campomanesia (Myrtaceae) from
Bahia, Brazil, based on specimens collected by J.S.
Blanchet over 1 50 years ago by Leslie R. Undrum and
Maru Ibrahim U. de Oliveira— 4(2):603

A new varietal combination, typification, and nomen-

termountain flora by Richard Spellenberg— 4(1):207

A new variety of Bromus flexuosus (Poaceae:
Pooideae: Bromeae: sect Bromopsis) by Ana Maria
Planchuelo— 4(2):653

Acronyms for recently proposed angiosperm families
by Daniel L Nickrent — 4(1):309

Additions to the vascular flora of New Mexico by Ben
S. Legler— 4(2):777

Agalinis flexicaulis sp. nov. (Orobanchaceae: Lami-
ales), a new species from northeast Florida by John
F Hays— 4(1 ):1

Annotated checklist of the vascular flora of the Jack
Gore Baygall and Neches Bottom Units of the Big
Thicket National Preserve, Tyler, Jasper, and Hardin
counties, Texas by Urry E. Brown, Barbara R. MacRob-
ERTs, Michael H. MacRoberts, Warren W. Pruess, I. Sandra
Elsik, and Stanley Jones— 4(1 ):473

Annotated checklist of the vascular plants of Fort Hood,
Texas by Uura L. Hansen— 4(1 ):523

WITH 164 Authors:

4(2010)

supporting Journal of the Botanical Research Institute of Texas.
Mississippi by Charles T. Bryson and Paul E. Rotnrock
— 4(1):347

Checklist of the vascular plants of Allegheny County,
Pennsylvania by Cynthia M. Morton and Loree Speedy
— 4(1):435

Coleataenia Griseb. (1879): the correct name for
Sorengia Zuloaga & Morrone (2010) (Poaceae:
Paniceae) by Robert J. Soreng— 4(2):691
Comments on a revision of Celtis subgenus Mertensia
(Celtidaceae) and the recognition of Celtis pallida
by James Henrickson— 4(1 ):287
Congruence between allometric coefficients and
phylogeny in stipoid grasses: an evo-devo study by
Jack Maze— 4(2):693

paraceae) and a key to the genera of neotropical
Capparaceae with variously stellate to peltate indu-
menta by Xavier Cornejo and Hugh H. Iltis— 4(1 ):3 1 1
Croton bigbendensis Turner (Euphorbiaceae) revis-
ited by James Henrickson — 4(1):295

Diversity and abundance of orchids in a Peruvian cloud
, Janovec, Eric
Christenson, John E. Pinder III, and Keri McNew Barfield
— 4(1):317

Diversity, natural history, and conservation of Vanilla
(Orchidaceae) in Amazonian wetlands of Madre de
Dios, Peru by Ethan Householder, John Janovec, Angel
Balarezo Mozambite, Javier Huinga Maceda, Jason Wells,
Renan Valega, Helena Maruenda, and Eric Christenson
-4(1):227

s (Asteraceae) density as a baseline
? climate change in La Sal Mountain
habitats by James F Fowler and Barb Smith— 4(2):747

First report of Persicaria hispida (Polygonaceae) from
North America north of Mexico (Texas) by Daniel E.
Atha and Wiluam Carr— 4(1 ):559
Floristic composition and potential competitors in
Lindera melissifolia (
sippi with reference t(

S. Hawkins, Daniel A. Skojac Jr., Nathan M. Schiff, and
Theodor D. Leininger-4(1):381
Flourensia ilicifolia (Compositae: Heliantheae),

789

distribudon para la especie by M. Socorro GonzAlez-

Quintanilla, JovAn AlemAn Medrano, Jaime Sanchez Salas

— 4(1):313

Folia taxonomica 17, Dilkea (Passifloraceae) 2. Con-
spectus of the species of the Guianas with three
new species by Christian Feuillet— 4(1):55
Folia taxonomica 1 8.The status of Passiflora citrifolia
and a new species in subgenus Astrophea (Pas-
sifloraceae), Passiflora jussieui by Christian Feuillet
— 4{2):609

Folia taxonomica 19.Typifications in Dilkea (Passiflo-
raceae) by Christian Feuillet — 4(2):615
Four nomendatural changes in Viola (Violaceae) by R.
John Little and Landon E. McKinney — 4{1):225

Hedychium forrestii (Zingiberaceae) with a new
synonymy and a variety from India by E. Sanoj, M.
Sabu, andT. Rajesh Kumar — 4(2):633

Listado floristico y aspectos ecologicos de la familia
Poaceae para Chihuahua, Durango y Zacatecas,
Mexico by Yolanda Herrera Arrieta y Armando CoRits
Ortiz— 4(2):71 1

Miliusa wayanadica (Annonaceae), a new species
from Western Ghats, India by M.K. Ratheesh Narayanan,
P. Sujanapal, N. Anil Kumar, N. Sasidharan, and M.
SiVADASAN— 4(1):63

Mirandea grisea (Acanthaceae), newforCoahuila and
Durango, Mexico by Eduardo Estrada-CastillOn, Jos6
Angel Villarreal-Quintanilla, and Jorge Arturo Alba- Avila

— 4(2):739

Rosa LOpez-Ferrari — 4(1):221
New names and combinations in the flora of Colorado.

XIII by W.A. Weber and R.C. Wittmann— 4(1 ):2 13
New vascular plant records for South Dakota by Gary
E. Larson— 4(1 ):467

Nomendatural transfers in the genus Myrsine (Myrsi-
naceae) for New Caledonia by Jon M. Rkzketson and
JohnJ. PipolyIII— 4(2):627

and Yunpeng Zhao — 4{ 1 ):28 1
Occurrence of anisophylly and anisoclady within
the Amaranthaceae by Donald B. Pratt and Lynn G.

1 . and sp. nov (Cyperaceae)
in Cretaceous Canadian amber by George Poinai^ Ja
AND David J. Rosen-4(2):685

Paspalum pubiflorum and P.quadrifarium (Poaceae)
new to California, with a key and notes on invasive

Travis Columbus — 4(2):761

Penstemon oklahomensis (Scrophulariaceae) in
Texas by Jeffrey N. Mink, Jason R. Singhurst, and Walter
C Holmes— 4(1 ):471

Phylogenetic analyses of the Gaylussacia frondosa
complex (Ericaceae: Vaccinieae) based on molecular
and morphological characters by Michael T. Gajoec-

zka, I<

Norris H, Williams, and Kent D. Perkins— 4(1 ):245
Planaltina nuevo genero de la tribu Spermacoceae
(Rubiaceae), endemico del planalto central de Brasil
y una nueva especie del Estado de Goias, Brasil by
Roberto M. Salas and Elsa L Cabral— 4(1): 193
Plant communities ofselectedembayments along the
mid- to mid-upper Ohio River floodplain by Joseph
S. Ely and Dan K. Evans— 4(1):41 1
Plants new to Florida by Richard R Wunderlin, Bruce F.
Hansen, Alan R. Franck, Keith A. Bradley, and John M.
Kunzer— 4(1):349

Poa ramifer (Poaceae: Pooideae: Poeae: Poinae), a
new aerially branching gynomonoecious species
from Peru by Robert J. Soreng and Paul M. Peterson
— 4(2):587

Poa unispiculata, a new gynodioecious species of
cushion grass from Peru with a single spikelet per
inflorescence (Poaceae: Pooideae: Poeae: Poinae)
by Gerrit Davidse, Robert J. Soreng, and Paul M. Peterson

-4(1):37

Polycarpon tetraphyllum (Caryophyllaceae)
new to the flora of Louisiana by Charles M. Al-
TIAN MeGYERI, and BrAD

tha subsp. pinnatisecta (Sacramento Prickly Poppy,
Papaveraceae) and re-evaluation of its taxonomic
status by Sandy D. Cervantes, Phil Tonne, Rajanikanth
Govindarajulu, Patrick J. Alexander, and C Donovan
Bailey— 4(l):261

Potentilla uliginosa (Rosaceae: Rosoideae), a new
presumed extinct species from Sonoma County,
California by Barry C. Johnston and Barbara Ertter

— 4(1):13

Clark— 4(1 ):271

790

Registro de dos nuevas localidades y reubicacion
de individuos de una de ellas de Astrophytum
myriostigma (Cactaceae) en Durango, Mexico by
Jaime Sanchez Salas, Gisela Muro P£rez, Eduardo Estrada
CastillOn y Mario GarcIa Aranda— 4(2):741
Rehabilitacibn y leaotipificacibn del gbnero Tessiera,

Spermacoceae): una nueva combinacion y un
nuevo sinonimo by Roberto M. Salas and Elsa L. Cabral
— 4(1):181

Revision of Lobelia sect. Plagiobotrys (Campanulace-
ae: Lobelioideae) by Thomas G. Lammers— 4(1):169
Revision of Lobelia sect. Speirema (Campanulaceae:
Lobelioideae) by Thomas G. Lammers— 4(1 ):1 59

Salvia coriana sp. nov. (Lamiaceae), a new species
from a cloud forest in western Guatemala by Taylor
Sultan Quedensley and Mario E. V6uz PErez— 4(1):27
Sarracenia minor var. okefenokeensis (Sarraceniace-

area by Jacob S. Thompson — 4{2):771
Scallopleaf sage (Salvia vaseyi: Lamiaceae) discovered
in Arizona by James W. Cain, III, Brian D. Jansen, Richard
S. Felger, and Paul R. Krausman — 4(2):755
Studies in Capparaceae XXVII: six newtaxa and a new
combination in Quadrella by Xavier Corner and Hugh
H. iLTis— 4(1):75

Studies in Cappai

raceae XXVIll: The Quadrella cyn-
complex by Hugh H. Iltis and Xavier

CoRNEX)— 4(1):93

Studies in the Capparaceae XXIX: synopsis of Qua-
drella, a Mesoamerican and West Indian genus by
Hugh H. Iltis and Xavier Cornex) — 4(1):117
Stylidium darwinii (Stylidiaceae), a new trigger plant
from Western Ghats of Karnataka, India by Sachin A.
Punekar and P. Lakshminarasimhan — 4(1):69
Supplemental notes on Bolivian Xyris (Xyridaceae) by
Robert Kral — 4(2):563

Taxonomic notes on the genera Stenotis and
Carterella (Rubiaceae) and transfer of Hedyotis
greenei to Stenotis by Edward E. Terrell and Harold
Robinson— 4{2):61 9

Ten new Myrtaceae from eastern and northeastern
Brazil by Marcos Sobral— 4{1):133
The Dominican amber fossil Lasiambix (Fabaceae:

aceae) by Kenton L Chambers and George 0. Ponar Jr.
-4(1):217

The ferns and lycophytes of a montane tropical forest
in southern Bahia, Brazil by Fernando B. Matos, AndrE
M. Amorim, and Paulo H. Labiak— 4(1):333
The genus Rytidosperma (Poaceae) in the United
States of America by Stephen J. Darbyshire, Henry E.
CoNNoa and Barbara Ertter — 4(2):663

and a new key to the genus by Mary Ann E. Feist

Kerr County, Texas by J
Hansen, Jeffrey N. f
and Walter C. Holmes— 4(1 ):49 7

The vascular plants of Mowotony Prairie: a small rem-
nant coastal grassland in Brazoria County, Texas by
DJ. Rosen— 4(1 ):489

Thomas Walter's species of Hedysarum (Leguminosae)
by Daniel B. Ward— 4(2):705
ThomasWalter's species of Melanthium (Liliaceae) by
Daniel B. Ward— 4(1 ):303

Transfer of Hedyotis intricata to Arcytophyllum

(Rubiaceae) by Edward E. Terrell and Harold Robinson
-4(2):625

Two new Andean species of Solanum section Crini-
tum (Solanaceae) by FrankT. Farruggia, Michael H. Nee,
AND Lynn Bohs— 4(2):595

Two new Bolivian species of Aulonemia (Poaceae:
Bambusoideae: Bambuseae) by Emmet J. Judziewicz,
Eddie L Shea, and Tanya M. Wayda— 4(2):569
Two new taxa of Scutellaria section Resinosa (Lami-
aceae) from northern Arizona by S.L Rhodes andTJ.
Ayers— 4(1 ):19

Typifications of names in Agalinis, Gerardia, and
Tomanthera (Orobanchaceae) by J.M.Canne-Hilliker
AND John F. Hays— 4(2):677

Una nueva especie de Fevillea (Cucurbitaceae: Zanon-
ieae) de Costa Rica by Armando Estrada Ch. and Daniel
SANTAMARiAA.— 4(1):45

Validation of Exochordeae (Rosaceae) by James L.
Reveal— 4(1 ):2 15

Vascular flora and edaphic characteristics of saline
prairies in Louisiana by Christopher S. Reid, Patricia L.
Faulknei^ Michael H. MacRoberts, Barbara R. MacRoberts,
AND Marc Bordelon— 4(1 ):357
Vascular flora of the Old Mulkey Meeting House State
Historic Site, Monroe County, Kentucky by Ralph L.
Thompson and Ronald L. Jones— 4(1 ):391

791

Index of 164 Authors:

Volume 4 ( 2010 )

Thank you for choosing JoumaJ of the Botanical Research Institute of Texas.

Alexander, Patrick J.— 4(1 ):261
Allen, Charles M.— 4(2):775
Amorim, Andre M.— 4{1):333
Anil Kumar, N.— 4(1 ):63
Armstrong, Bill— 4(1 ):497
Arturo Alba-Avila, Jorge
— 4(2):739

Atha, Daniel E.— 4(1):559
Ayers, TJ.—4(1):19

Bailey, C Donovan— 4(1 ):261
Balarezo Mozambite, Angel
— 4(1):227

Barfield, Keri McNew— 4(1):317
Barringer, Kerry— 4(1 ):51
Bohs, Lynn— 4(2):595
Bordelon, Marc— 4(1 ):357
Boufford, David E.— 4(1):281
Bradley, Keith A.— 4(1):349
Brown, Larry E.— 4(1):473
Bryson, Charles T— 4(1 ):347

Cabral, Elsa L— 4(1):181, 193
Cain, III, James W.— 4(2):755
Canne-Hilliker, J.M.— 4(2):677
Carr, William— 4(1 ):559
Cervantes, Sandy D.— 4(1):261
Chambers, Kenton L— 4(1):217
Christenson, Eric— 4(1 ):227, 317
Clark, Lynn G.— 4(1 ):271
Columbus, J. Travis— 4(2):761
Connor, Henry E.— 4(2):663
Cornejo, Xavier— 4(1 ):75, 93.

Cortes Ortiz, Armando— 4(2):71 1

Derbyshire, Stephen J. — 4(2):663
Davidse, Gerrit— 4(1):37
Denham, Silvia 5.— 4(2):761

Elsik,l. Sandra— 4(1 ):473
Elven, Reidar— 4(1):219
Ely, Joseph S.— 4(1):411
Ertter, Barbara— 4(1 ):13; 4(2):663
Espejo-Serna, Adolfo— 4(1 ):221

Estrada Castillon, Eduardo— 4(2):
739, 741

Estrada Ch„ Armando— 4(1 ):45
Evans, Dan K.— 4(1):411

Farruggia, Frank T.—4(2):595
Faulkner, Patricia L— 4(1):357
Feist, Mary Ann E.— 4(2):641
Felger, Richard S.— 4(2):755
Feuillet, Christian— 4(1 ):55;

4(2):609;4(2):615
Fowler, James F— 4(2):747
Franck, Alan R — 4(1):349
FreIsJr., Donnie— 4(1 ):497
Fu, Chengxin— 4(1):281

Gajdeczka, Michael!.— 4(1 ):245
Gandhi, Kanchi N.— 4(1):219
Garcia Aranda, Mario— 4(2):741
Gonzalez-Elizondo, M. Socorro
— 4(1):313

Govindarajulu, Rajanikanth
— 4(1):261

Grandon, Jarrod— 4(2):775

Hansen, Bruce F— 4(1):349
Hansen, Laura L— 4(1):497, 323
Hawkins, Tracy S.— 4(1):381
Hays, John F— 4(1):1; 4(2):677
Helliwell, Richard— 4(1 ):7
Henrickson, James— 4(1 ):287,
295; 4(2):581
Herrera Arrieta, Yolanda
— 4(2):71 1

Holmes, Walter C— 4(1 ):471, 497
Householder, Ethan— 4(1 ):227
Huinga Maceda, Javier — 4(1):227

Ibrahim U. de Oliveira, Marla
— 4(2):603

litis, Hugh H.— 4(1 ):75, 93. 117,
311

Janovec, John R— 4(1):227, 317
Jansen, Brian D.— 4(2):755
Johnston, Barry C.— 4(1):13

Jones, Ronald L— 4(1):391
Jones, Stanley— 4(1 ):473
Judd, Walters.— 4(1 ):245
Judziewicz, Emmet J.— 4(2):569

Krai, Robert— 4(2):563
Krausman, Paul R.— 4(2):755
Kumar,! Rajesh— 4(2):633
Kunzer, John M.— 4(1 ):349

Labiak, Paulo H.— 4(1 ):333
Lakshminarasimhan, P— 4(1):69
Lammers, Thomas G.— 4(1 ):1 59,
169

Landrum, Leslie R.— 4(2):603
Larson, Gary E.— 4(1 ):467
Legler, Ben S.— 4(2):777
Leininger, Theodor D.— 4(1):381
Little, R. John— 4(1 ):225
Lopez-Enriquez, I. Lorena
-4(1):313

Lbpez-Ferrari, Ana Rosa— 4(1):221
Luke, Rebecca Repasky— 4(1):317

MacRoberts, Barbara R.— 4(1):
357, 473

MacRoberts, Michael H. — 4(1):
357, 473
Martinez-Correa,

Nancy^l):221
Maruenda, Helena — 4(1):227
Matos, Fernando B.^(l):333
Maze, Jack— 4(2):693
McKinney, Landon E.— 4(1):225
Medrano, Jovan Aleman— 4(1):
313

Megyeri, Krisztian^2):775
Mink, Jeffrey N.— 4(1):471, 497
Morton, Cynthia M. — 4(1):435
MuroPere4Gisela— 4(2):741
Murray, David F— 4(1):219

Nee, Michael H.— 4(2):595
Neubig,KurtM.— 4(1):245
Nickrent, Daniel L— 4(1):309

792

Journal of the Botanical Research Institute of Texas 4(2)

Perkins, Kent D.— 4(1):245
Peterson, Paul M.— 4(1):37; 4(2):
587

Pinder III, John E.— 4(1):317
PipolyllUohnl— 4(2):627
Planchuelo, Ana Maria— 4(2):653
Poinar Jr., George 0.-4(l):217;
4(2):685

Pratt, Donald B.— 4(1):271
Pruess, Warren W.— 4(1 ):473
Punekar, Sachin A.— 4(1):69
Puppo, Pamela— 4(1 ):33

Quedensley,Taylor Sultan— 4(1 ):27

Ratheesh Narayanan, M.K.— 4(1 ):63
Reid, Christopher S.— 4(1):357
Reveal, James L—4(l):215
Rhodes, S.L— 4(1 ):19
Ricketson, Jon M.— 4(2):627
Riefner, Jr., Richard E.— 4(2):761
Robinson, Harold— 4(2):61 9, 625
Rosen, DJ.— 4(1):489; 4(2):685
Rothrock, Paul E.— 4(1):347

Sabu, M.— 4(2):633
Salas, Roberto M.-4(l):181. 193
Sanchez Salas, Jaime— 4(1):313;
4(2):741

Sanoj, E.— 4{2):633
Santamaria A., Daniel — 4(1):45
Sasidharan, N. — 4(1):63
Schiff, Nathan M.— 4(1):381
Shea, Eddie L— 4(2):569
Singhurst, Jason R.-4(1):471,
497

Sivadasan, M.— 4(1):63
Skojac Jr., Daniel A.-4(l):381
Smith, Barb— 4(2):747
Sobral, Marcos— 4(1 ):133
Soreng, Robert J,— 4(1):37;

4{2):587,691
Speedy, Loree-4(1):435
Spellenberg, Richard— 4(1 ):207
Sujanapal, R— 4(1):63

Terrell, Edward E.— 4(2):619, 625
Thompson, Jacob S.— 4(2):771

Thompson, Ralph L— 4(1):391
Tonne, Phil— 4(1 ):261

Valega, Renan— 4(1 ):227
Van Devender, Thomas R.— 4(2):
581

Veliz Perez, Mario E.— 4(1):27
Villarreal Quintanilla, Jose A.— 4(1):
313; 4(2):739

Waguespack, Brad-4(2):775
Ward, Daniel B.— 4(2):303, 705
Wayda, Tanya M.— 4(2):569
Weber, W.A.— 4(1 ):2 13
Wells, Jason— 4(1 ):227
Whitten, W. Mark— 4(1 ):245
Williams, Norris H.— 4(1):245
Wipff, III, Joseph K.— 4(2):683
Wittmann, R.C— 4(1):213
Wunderlin, Richard R— 4(1):349

Xiang, Qiu-Yun (Jenny)— 4(1 ):281
Zhao, Yunpeng— 4(1 ):281

Botanical Names and Subject Index:

Volume 4 (2010)

New names (162) in bold face

bakeri-4(l):209
crux-maltae— 4(1):208
robusta— 4(1):209
Acanthaceae— 4(2):739
Achnatherum

hendersonii— 4(2):696
lemmonii— 4(2):696
nelsonii— 4(2):696
occidentale— 4(2):696

divaricata— 4(1):5
filifolia— 4(1):5
flexicaulls— 4(1):1,5

linifotia— 4(1):5
iongifolia— 4(2):677

maritima var. gracilis— 4(2):677
maritima var. pumila— 4(2):678
microphylla— 4(2):678
obtusifolia— 4(1):5;4(2):678
palustris var. corymbosa— 4(2):
678

palustris var. divaricata — 4(2):
678

plunkenetii— 4(1):5
setacea var. humilis— 4(2):678
setacea var. major— 4(2):678
tenuifolia-4(l):5
tenuifolia var. humilis— 4(2): 679
tenuifolia var. saxatilis— 4(2):679
virgata— 4(2):679
Alabama— 4(1 ):347
Allegheny County — 4(1 ):435
Allionia

glandulifera— 4(1):209
incarnata var. glabra— 4(1 ):209

Allometric coefficients — 4(2):693
Alternanthera— 4(1 ):278
Amaranthaceae— 4(1):271
Amazonian wetlands— 4(1 ):227
Amber— 4(2):685

4(1):305

Amphorogynaceae— 4(1 ):309
Andean— 4(2):595
Angelonia

angustifolia— 4(1):53
biflora— 4(1 ):53
ciliaris— 4(1):53
gardneri— 4(1):53
parviflora— 4(1):51, 53
pilosella— 4(1):54
Angiosperm families— 4(1 ):309
Anisoclady— 4(1):271
Anisophylly — 4(1 ):271
Annonaceae — 4(1 ):63

793

Anthospermopsis— 4(1 ):1 95
Apera interrupta — 4(1):467
Apiaceae— 4(1):7; 4(2):641
Araucaria heterophylla— 4{1):349
Arcytophyllum— 4(2):625
fasciculatum— 4(2):625
Ardisia solanacea— 4(1):350
Argemone

pinnatisecta— 4(1):268

pleiacantha subsp. pinnatisecta
— 4(1):261

Arizona— 4(1 ):1 9; 4(2):755
Artemisia ludoviciana— 4(1):350
Asteraceae^d ):3 1 3; 4(2):747
Astrophea (subg.)— 4(2):609
Astrophytum myriostigma — 4(2):
741

Aulonemia
boliviana— 4(2):570

herzogiana^ — 4(2):570
longipedicellata— 4(2):570
scripta— 4(2):573
tremula— 4(2):570
Azolla mexicana— 4(1):468

Bahia— 4(1 ):333; 4(2):603
Bambuseae— 4(2):569
Bambusoideae— 4(2):569
Beautempsia — 4(1):31 1
Big Thicket National Preserve—
4(1):473

Blanchet,J.S.— 4(2):603
Blutaparon— 4(1):278
Boerhavia spicata var. torreyana—
4(1):208

Bolivia— 4(2):563, 569
Botrychium
hesperium— 4(2):779

minganense— 4(2):780
multifidum— 4(2):780
pinnatum— 4(2):780
tunux— 4(2):781

Brasil— 4(1 ):1 33, 193, 333;4(2):603
Brazoria County— 4(1 ):489
Bromeae— 4(2):653
Bromeliaceae— 4(1):221

Bromopsis (Sect.)— 4(2):653
Bromus

flexuosus— 4(2):653
4(2):655

flexuosus var. famatinensis—
4(2):657

flexuosus var. flexuosus— 4(2):
655

lanatus — 4(2):655
modestus— 4(2):655
squarrosus— 4(2):781
Buckleya— 4(1):281
angulosa— 4(1):285
distichophylla— 4(1 ):285
graebneriana— 4(1):285
henryi— 4(1):285
lanceolata— 4(1):285
Butia capitata— 4(1):350

tala— 4(1 ):291
Cervatesiaceae— 4(1 ):309
Chamaelirium luteum — 4(1):305
Chihuahua— 4(2):711
China— 4(1 ):281

Cinnamomum verum— 4(1):350
Cloud Forest— 4(1 ):27, 317
Coahuila— 4(2):739

anceps subsp. rhizomata—
4(2):691

longifolia subsp. abscissa—
4(2):691

Cactaceae— 4(2):741

Calceolariaceae— 4(1 ):33
California— 4(1 ):1 3; 4(2):761
Campanulaceae— 4(1):159, 169

eugenioides— 4(2):604
Canada— 4(2):685
Capparaceae^(1):75,93, 1 17,31 1

leavenworthii— 4(1 ):468
microglochin— 4(2):778
nelsonii— 4(2):778
oklahomensis — 4(1):347
pseudocyperus— 4(1 ):468
Carterella— 4(2):619

Celastrus paniculatus— 4(1):350
Celtidaceae— 4(1):287
Celtis

ehrenbergiana — 4(1):289
iguanaea— 4(1):288
subg. Mertensia-4(1):287
pallida— 4(1 ):287

pallid var. discolor— 4(1 ):292
pallid var. pallida— 4(1):292

4(2):691

longifolia subsp. rigidula—
4(2):691

petersonii— 4(2):692
prionitis — 4(2):692
stenodes— 4(2):692
tenera— 4(2):692

Colorado— 4(1 ):2 13
Comandraceae — 4(1):309
Compositae — 4(1):31 3
Congruence — 4(2):693
Conyza laevigata— 4(1 ):350
Cordia curassavica— 4{1):350
Corrigenda — 4(1):31 1
Costa Rica— 4(1 ):45
Coutaceae— 4(1):309
Crataegus chrysocarpa var.

chrysocarpa— 4(2):782
Cretaceous Canadian amber—
4(2):685

Crinitum (Sect.)— 4(2):595
Croton bigbendensis — 4(1):295
Cucurbitaceae— 4(1 ):45
Curculigo capitulata— 4(1):351
Cushion Grass— 4(1):37
Cydista aequinoctialis — 4(l)d51
Cyperaceae^1):347; 4(2):685
Cyperus unioloides— 4(1)351

794

Desmodium
cuspidatum — 4(2):708
glutinosum— 4(2):707
hirta— 4{2):708
laevigatum — 4(2):707

Euphorbiaceae— 4(1):295
Evo-Devo— 4(2):693

Exochordeae— 4(1):215

Gomphrena— 4(1):278
Gossypianthus— 4(1 ):278
Grewia asiatica— 4(1):352
Guatemala (Western)— 4(1 ):27
Guianas— 4(1):55,4(2):609

nudiflorum— 4(2):709
paniculatum— 4(2):709
repens— 4(2):708
strictum— 4(2):707
stuevei— 4(2):708
tenuifolium— 4(2):707

Fabaceae— 4(1):217
Ferns— 4(1 ):333
Festuca hallii— 4(2):781
Fevillea narae— 4(1):45
Flora Caroliniana— 4(2):705
Florida— 4(1 ):1, 349

Dieffenbachia seguine— 4(1):351
Dilkea— 4(2):615
clarkei— 4(1):56,61
subg. Dilkea— 4(1 ):61
subg.Epkia-4(l):59
ercta— 4(1):61
exilis— 4(1):59,61

helleborifolia— 4(2):615
johannesii— 4(2):615
lecta— 4(1):61
retusa— 4(1):59
vanessae— 4(1):61
Dimocarpus longan— 4(1):351
Dominican amber— 4(1 ):2 17
Draba streptobrachia— 4(2):778
Drynaria quercifolia — 4(1):351
Durango— 4(1 ):3 13; 4(2):711,
739, 741

Ecuador— 4(1 ):33
Edaphic— 4(1):357
Embayments— 4(1):41 1
Ericaceae— 4(1 ):245
Erigeron mancus— 4(2):747
Eriochloa

acuminata— 4(1 ):351
villosa— 4(1):468
Eriogonum arcuatum var.

xanthum— 4(2):782
Eriophorum scheuchzeri— 4(2):
778

Eucalyptus camaldulensis — 4(1):
352

Eugenia

cernua— 4(1):314
dentata — 4(1):314
ilicifolia— 4(1):313,314
microphylla— 4(1):315
monticola-4(1):315
pringlei — 4(1):315
pulcherrima— 4(1):315
retinophylla— 4(1):314
solitaria— 4(1):314
Fort Hood Texas— 4(1 ):523
French Guiana 4(2):609
Froelichia— 4(1):278

Galium circaezans — 4(1):468
Gastrolychnis hitchgulrei— 4(1):
213

Gaylussacia frondosa— 4(1 ):245
Georgia— 4(1 ):347
Gerardia— 4(2):677
aspera— 4(2):680
filifolia— 4(2):680

maritima— 4(2):679
purpurea var. crassifolia— 4(2):

purpurea var. parviflora — 4(2):

purpurea var. paupercula —

4(2):680

tenuifolia var. leptophylla—

4(2):680

tenuifolia var. macrophylla —
4(2):681

Glaux maritima— 4(1 ):468
Goias— 4(1):193

Hardin County-4(1):473
Hechtia podantha— 4(1):222
Hedychium

forrestii var. forrestii— 4(2):633,
634

forrestii var. latebracteatum—
4(2):634

forrestii var. pala-
niense— 4(2):634
Hedyotis— 4(2):619
greenei— 4(2):619
intricata— 4(2):625
Hedysarum— 4(2):705
Heliantheae— 4(1):313
Helonias bullata— 4(1):305
Heterotheca pumila— 4(2):777
Houttuynia cordata— 4(1):352

India— 4(1 ):63, 69; 4(2):633
Intermountain flora— 4(1 ):207
Invasive species— 4(2):761
Iresine— 4(1):278

Jack Gore Baygall Unit— 4(1):473
Jasper County — 4(1):473

alpinoarticulatus— 4(2):778
biglumis— 4(2):778
parryi— 4(2):778
triglumis var. triglumis — 4(2):
779

Karnataka— 4(1 ):69
Kentucky— 4(1 ):391
Kerr County— 4(1 ):497
Kerr Wildlife Management Area —

4(1):497

Kobresia simpliciuscula— 4(2):778

La Sal Mountains— 4(2):747
Lagotis

glauca subsp. glauca — 4(1 ):2 1 9

glauca subsp. lanceolata^
4(1):220

Lamiaceae— 4(1 ):1 9, 27; 4{2):755
Lamiales — 4(1 ):1
Lauraceae— 4(1):381
Lechea stricta— 4(1):468
Leguminosae— 4(2):705
Leucophyllum
hintonorum— 4(2):584

Liliaceae— 4(1):303
Lindera melissifolia— 4(1):381
Lipocarpha micrantha— 4(1):468
Listera borealis— 4(2):781

borneensis — 4(1 ):1 73
borneensis subsp. borneensis
— 4(1):173

borneensis subsp. celebensis
— 4(1):174

borneensis subsp. grandiflora
— 4(1):176

-4(1):174

fangiana— 4(1):162

montana— 4(1):162
origenes— 4(1):177
sect. Plagiobotrys— 4(1):172

Lobelioideae— 4{1):159, 169

perenne— 4(2):683
perenne subsp. stoloniferum
— 4(2):683
Lomatium

ambiguum— 4(1):11
canbyi— 4(1):10
cous-4(1):11

hambleniae— 4(1):11
henersonii— 4(1):11
leptocarpum— 4(1):1 1

roseanum — 4(1 ):1 1
watsonii-4(1):11
Louisiana— 4(1 ):347, 357; 4(2):775
Lumnitzera racemosa— 4(1):352
Lycophytes— 4(1):333

Madrede Dios— 4(1 ):227
Melanthium— 4(1):303
virginicum— 4(1):305
Mesoamerica— 4(1 ):1 1 7
Mexico— 4(1 ):5 1,313; 4(2):581 ,
711,739,741
Microteaceae— 4(1 ):309
Miliusa wayanadica— 4(1):64
Mimusops coriacea— 4(1):352
Mirandea grisea— 4(2):739
Mississippi— 4(1 ):347, 381
Mitracarpus— 4(1):195
Monotropa uniflora— 4(1 ):468
Mowotony Prairie— 4(1 ):489
Myrcia

tetraphylla- 4(1):152

Myrsinaceae— 4(2):627
Myrsine— 4(2):627

arborea— 4(2):628
asymmetrica — 4(2):628

asymmetrica subsp. magnifolia
— 4(2):628

lanceolata subsp. ouena-

is— 4(2):630
modesta— 4(2):630

modesta subsp. corlaria—
4(2):630

novocaledonica subsp. boulin-
daensis— 4(2):630
novocaledonica subsp. kaa-
laensls— 4(2):630

novocaledonica subsp. plro-
guensis— 4(2):630
oblanceolata— 4(2):630

parvicarpa subsp. amossensis
— 4(2):631

parvicarpa subsp. pachyphylla
-4(2):631

pronyensis— 4(2):631

piperi-4(1):11
rollinsii— 4(1):11

tchingouensis— 4(2):63 1
verrucosa— 4(2):631

verrucosa subsp. microphylla
— 4(2):631
yateensis— 4(2):632
Myrtaceae— 4(1):133
Myrtaceae— 4(2):603

Najas minor— 4(1 ):469
Nanodeaceae— 4(1):309
Neches Bottom Unit— 4(1 ):473
New Caledonia — 4(2):627
New Mexico— 4(2):777
Nuttallia rhizomata — 4(1):213
Nyctaginaceae — 4(1 ):207

Ochoco Mountains— 4(1 ):5
Ohio— 4(1 ):41 1
Ohio River— 4(1 ):411
Okefenokee Swamp Area— 4(2):
771

Old Mulkey Meeting House State
Historic Site— 4(1 ):391
Orchidaceae— 4(1):227,317
Oregon— 4(1 ):5

Oreocarya revealii— 4(1):213

Orobanchaceae— 4(1):565; 4(2):
677

Oxybaphus glaber— 4(1):208

Papaveraceae— 4(1 ):261
Paramos— 4(1 ):33
Paspalum— 4(2):761
dilatatum— 4(2):767
distichum— 4(2):767
notatum— 4(2):767
notatum var. saurae— 4(2):767
pubiflorum— 4(2):761,767
quadrifarum— 4(2):761, 767
urvillei— 4(2):767
vaginatum— 4(2):767
Passiflora— 4(2):609
cardonae — 4(2):61 3
cauliflora— 4(2):613

cerradensis— 4(2):613
citrifolia— 4(2):609
jussieui— 4(2):611,613
maguirei— 4(2):613
skiantha— 4(2):613
Passifloraceae— 4(1 ):55; 4(2):609,
615

Pennsylvania— 4(1 ):435
Penstemon oklahomensis— 4(1):

Pentas lanceolata — 4(1):353
Persicaria hispida— 4(1):559
Peru— 4(1 ):37, 227, 317;4(2):587
Pfaffia— 4(1):278
Philodendron hederaceum —
4(1):353

Pimentadioica— 4(1):353
Pithecellobium bahamense—
4(1):353

Planaltina— 4(1):193, 195, 198
capitata— 4(1):199
lanigera— 4(1):202

nuttallii— 4(2):648
texense— 4(2):641,648
Purshia tridentata— 4(1):469

Quadrella

alaineana— 4(1):75, 129
angustifolla— 4(1):129
antonensis— 4(1):125
asperifolia— 4(1):125,126

calciphila— 4(1):125,126

cynophallophora— 4(1 ):93
cynophallophora f. iinearifolia
— 4(1):98, 122

dressleri— 4(1):77,125
ferruginea— 4(1):130

Plant communities— 4(1 ):41 1
Plantaginaceae — 4(1):51, 219
Plinia espinhacensis— 4(1):156
Poa

ramifer— 4(2):587
unispiculata— 4(1):37

Poaceae (checklist)— 4(2):71 1
Poaceae— 4(1):37; 4{2):569, 587,
653,663,683,691,761
Poeae— 4(1):37; 4(2):587, 683
Poinae— 4(1):37, 587
Polemonium occidentale var.

occidentale— 4{2):782
Polyalthia suberosa— 4(1):353
Polycarpon tetraphyllum— 4(2):
775

Polygonaceae— 4(1 ):559
Pooideae— 4(1):37; 4(2):587, 653
Potentilla
nivea— 4(2):782

uliginosa— 4(1):14

Ptilagrostis ported— 4(2):781
Ptilimnium

capillaceum— 4(2):648
costatum— 4(2):646, 648

4(1):129,130

ferruginea subsp. ferruginea—
4(1):129

indica— 4(1):125,126
subg. intutis— 4(1):121, 128
isthmensis— 4(1):107, 122

4(1):110,1123

jamaicensis— 4(1):98, 121, 123
jamaicensisf longifolia— 4(1):

lindeniana— 4(1):83, 125
lundellii— 4(1):125,127
mirifica— 4(1):125, 127
morenoi— 4(1):86, 125

797

morenoi f. hastata— 4(1):90
odoratissima— 4(1):123
pringlei— 4(1):125
pringlei— 4(1):127
subg.Quadrella— 4(1):121
quintanarooensis— 4(1 ):1 04,
121

siliquosa— 4(1):98, 121
singularis— 4(1):129,131
steyermarkii— 4{1):125, 128

Radermachera sinica— 4(1):353
Rosa canina— 4(2):782
Rosaceae— 4(1):13,215
Rousselia humilis— 4(1):353
Rubiaceae— 4(1):181, 193; 4(2):
619,625

Rytidosperma— 4(2):663
biannulare— 4(2):665
caespitosum— 4(2):665
penidllatum— 4(2):665
racemosum — 4(2):665
richardsonii— 4(2):665

Sacramento Prickly Poppy— 4(1):
261

Saline prairies— 4(1 ):357

coriana-4(1):27

curtiflora— 4(1):28
excelsa— 4(1):28
hispanica— 4(1):28
holwayi— 4(1):28
polystachya — 4(1 ):28
purpurea — 4(1):28
tiliaeifolia— 4(1):28
vaseyi— 4(2):755
Santalaceae— 4(1):281
Sarracenia
minor— 4(2):771
minor var. okefenokeensis —
4(2):771

Sarraceniaceae— 4(2):771
Scallopleafsage— 4(2):755
Scrophulariaceae— 4(1 ):471 ; 4(2):
581

platyphylla— 4(1):24
platyphylla var. grahamiana—
4(1):26

platyphylla var. kaibabensis —

4(1):26

4(1):24

platyphylla var. occidentalis—
4(1):24

platyphylla var. platyphylla—
4(1):24

platyphylla var. tessellata—
4(1):26

potosina— 4(1):24
potosina var. kaibabensis—
4(1):20

Senna atomaria— 4(1):354
Solanaceae— 4(2):595
Solanum

umbellatum— 4(1):354
Sonoma Couinty— 4(1 ):1 3

South Dakota— 4(1 ):467
Spermacoceae— 4(1):181, 193
Staelia— 4(1):195
State Records
Arizona— 4(2):755
California— 4(2):761
Louisiana— 4(2):775
New Mexico— 4(2):755
South Dakota— 4(1 ):467
Texas— 4(1 ):471, 559
Stellaria parva— 4(1):354
Stenotis — 4(2):619
arenaria— 4(2):620
asperuloideds— 4(2):620
australis — 4(2):620
brevipes— 4(2):620
gracilenta— 4(2):621
greenei — 4(2):622
mucronata — 4(2):620
Stipoid grasses— 4(2):693
Strombosiaceae-4(1):309
Sty lid iaceae — 4( 1 ):69
Stylidium

darwinii— 4(1):69, 72, 75
kunthii— 4(1):73
tenellum— 4(1):73

Tessiera— 4(1):195

lanigera— 4(1):191

pubescens— 4(1):190
Texas— 4(1 ):471, 473, 489, 497,
523, 559

Thesiaceae— 4(1 ):281, 309
Thomas Walter— 4(1 ):303
Tidestromia— 4(1):278
Tofieldia racemosa— 4(1):305
Tomanthera lanceola-

Trema orientalis— 4(1):34
Trichosanthes cucumeri-
na-4(1):354
Triplaris melaenoden-
dron— 4(1):354
Tripterocalyx carneus var. p
culatus— 4(1):207
Tyler County— 4(1 ):473

United States of Ameri-
ca— 4(2):663
Utah— 4(2):747

Vaccinieae— 4(1):245
Vanilla— 4(1 ):227
bicolor— 4(1 ):230
cristato-callosa— 4(1):230
guianensis— 4(1):230
palmarum— 4(1):230
pompon subsp. grandiflo-
ra-4(1):230
riberoi— 4(1):230
Viola

epipsila var. repens— 4(1 )*,225
pedatifida var. brittoni-
ana— 4(1):225
praemorsa var. fiavovi-
rens— 4(1)*^25
sororia var. grisea— 4(1):226
Violaceae— 4(1):225
Walter, Thomas— 4(2):705

Scutellaria

798

Westindies— 4(1):117
Western Ghats— 4(1 ):63, 69
Wissadula amplissima— 4(1):355

Ximeniaceae— 4(1 ):309

Xyridaceae— 4(2):563
Xyris

crassifunda— 4(2):563

vacillans— 4(2):566
Zacatecas— 4(2):71 1
Zanonieae— 4(1):45
Zigadenus glaberrimus— 4(1 ):305
Zingiberaceae— 4(2):633

Agalinisflexicaulis Hays, sp. nov. — 4(1 ):1
Angelonia parviflora Barringer, sp. nov— 4(1):51
Arcytophyllum fasciculatum (A. Gray) Terrell & I

Lagotis glauca subsp. lanceolata (Hulten) D.F. Murray &
Elven stat. nov.— 4(1):220

Leucopyllum mojinense Henrickson & T.R. Van Devender,

s Lammers, subsp. nov.—

nov.— 4(2):691

caricoides (Nees exTrin.) Soreng, comb, nov.— 4(2):691
longifolia (Torr.) Soreng, comb, nov.— 4(2):691
longifolia subsp. abscissa (Swallen) Soreng, comb.

longifolia subsp. combsii (Scribner & C.R. Ball) Soreng,
comb, nov.— 4{2):691

longifolia subsp. elongata (Scribn.) Soreng, comb,
nov.— 4(2):691

longifolia subsp. rigidula (Bose ex Nees) Soreng, comb,
nov.— 4(2):691

petersonii (Hitchc. & Ekman) Soreng, comb, nov.—
4(2):692

prionitis (Nees) Soreng, comb, nov.— 4(2):692
stenodes (Griseb.) Soreng, comb. nov. — 4(2):692
tenera (Beyr. exTrin.) Soreng, comb, nov.— 4(2):692

clarkei Feuillet, sp. nov.-4(l):56
exilis Feuillet, sp. nov.-4{1):59
granvillei Feuillet, sp. nov. — 4(1):56

azeda Sobral, sp. nov.— 4(1 );1 33
valsuganana Sobral, sp. nov. — 4{1):136
xochordeae Schulze-Mentz ex Reveal,
4(1):215

nov.-4(1):174

origenes Lammers, sp. nov.— 4{1):1 77

sect. Plagiobotrys Lammers, sect, nov.— 4(1):172

4(1):161

[>lium perenne subsp. stoloniferum (C. Lawson) Wipff,
comb, et stat. nov. — 4(2):683

Miliusa wayanadica Sujanapal, Ratheesh & Sasidhara

crassa Sobral, sp. nov.— 4{1):1 38
floridissima Sobral, sp. nov.— 4(1 ):1 40
mucugensis Sobral, sp. nov.— 4(1):142
pendula Sobral, sp. nov.— 4(1):145

pseudospectabilis Sobral, sp. nov.— 4(1):1 50
tetraphylla Sobral, sp. nov.— 4(1):1 52

. Schmid) Ricketson & Pi

»v.— 4(2):628

jorea (M. Schmid) Ricketson & Pipoly, c

799

(Mez) R

4(2):628

asymmeti

&Pipoly,c

i Pipoly, c

& Pipoly, c

a (M. Schmid) Ricketson
nb.etstat. nov.-4(2):628
asymmetrica subsp. paniensis (M. Schmid) Ricketson &
Pipoly, comb, et stat. nov.— 4(2):628
asymmetrica subsp. parvifolia (M. Schmid) Ricketson &
Pipoly, comb, et stat. nov.— 4{2):628
. Schmid) Rickel

(M. Schmid) Rick

) Ricketson & Pipoly, comb, nov—

Schmid) Ricketson & Pipoly, comb.
Schmid) Ricketson & Pipoly, comb.

nov— 4(2):629

grandifolia (S. Moore) Ricketson & Pipoly, comb, nov—

4(2):629

humboldtensis (M. Schmid) Ricketson & Pipoly, comb,
nov— 4(2):629

katrikouensis (M. Schmid) Ricketson & Pipoly, comb.
nov-4(2):629

koghiensis (M. Schmid) Ricketson & Pipoly, comb.

kuebiniensis (M. Schmid) Ricketson & Pipoly, c

lecardii (Mez) Ricketson & Pipoly, comb, nov— 4(2):629
macrophylla (Mez) Ricketson & Pipoly subsp. me-
naziensis (M. Schmid) Ricketson & Pipoly, comb, et stat.

nov— 4(2):630

memaoyaensis (M. Schmid) Ricketson & Pipoly, comb,
nov— 4(2):630

4(2):630 ^

modesta subsp. coriaria (M. Schmid) Ricketson & Pipoly,

comb, et stat. nov— 4(2):630

d. Schmid) Ricketson &
t. nov— 4(2):630

I. Schmid) Ricketson & Pipoly, comb.

nov— 4(2):630

nigricans (M. Schmid) Ricketson & Pipoly, comb, nov—
4(2):630

novocaledonica subsp. balabioensis (M. Schmid) Rick-
etson & Pipoly, comb, et stat. nov— 4(2):630
novocaledonica subsp. boulindaensis (M. Schmid)
Ricketson & Pipoly, comb, et stat. nov— 4{2):630
novocaledonica subsp. kaalaensis (M. Schmid) Ricket-
son & Pipoly, comb, et stat. nov— 4(2):630
novocaledonica subsp. mueoensis (M. Schmid) Ricket-
son & Pipoly, comb, et stat. nov— 4(2):630

a subsp. piroguensis (M. Schmid) Rick-
etson & Pipoly, comb, et stat. nov— 4(2):630
oblanceolata (M. Schmid) Ricketson & Pipoly, comb,
nov— 4(2):630

oblanceolata subsp. doensis (M. Schmid) Ricketson &
Pipoly, comb, et stat. nov— 4(2):631
obovalifolia (M. Schmid) Ricketson & Pipoly, comb,
nov— 4(2):631

ouameniensis (M. Schmid) Ricketson & Pipoly, comb,
nov— 4(2):631

ouazangouensis (M. Schmid) Ricketson & Pipoly, comb,
nov— 4(2):631

ovicarpa (M. Schmid) Ricketson & Pipoly, comb, nov—

paniensis (M. Schmid) Ricketson & Pipoly, comb, nov—

4(2):631

parvicarpa (M. Schmid) Ricketson & Pipoly, comb,
nov— 4(2):631

parvicarpa subsp. amossensis (M. Schmid) Ricketson &

Pipoly, comb, et stat. nov— 4{2):631

parvicarpa subsp. pachyphylla (M. Schmid) Ricketson &

Pipoly, comb, et stat. nov— 4(2):631

poumensis (M. Schmid) Ricketson & Pipoly, comb.

nov— 4(2):631

pronyensis (Guillaumin) Ricketson & Pipoly, comb.

v.-4(2):631

>noph

4(2):631

, Schmid) Picket;
dez) Ricketson & Pipoly,
. Schmid) Ricketson &

Pipoly,

nov— 4(2):631

tchingouensis (M. Schmid) Ricketson & Rpoly, comb,
nov— 4(2):631

verrucosa (M. Schmid) Ricketson & Pipoly, comb, nov—
4(2):631

verrucosa subsp. microphylla (M. Schmid) Ricketson &

Pipoly, comb, et stat. nov — 4(2):631

yateensis (M. Schmid) Ricketson & Pipoly, comb, nov—

4(2):632

Oreocarya revealii

Journal of the Botanical Research Institute of Texas 4(2)

Planaltina R.M. Salas & E.L Cabral, gen. nov.— 4(t):198
capitata (K. Schum.) R.M. Salas & E.L Cabral, comb,
nov.— 4(1):199

lanigera (DC.) R.M. Salas & E.L. Cabral, comb, nov.—
4(1):202

myndeliana R.M. Salas & E.L Cabral, sp. nov.— 4(1):204
Plinia espinhacensisSobral, sp. nov.^(l):156
Poa

ramifer Soreng & P.M. Peterson, sf

v.-4(2):587

alaineana Cornejo & litis, sp. nov. — 4
4(?):129

antonensis (Woodson) litis & Corn
4(1):125

asperifolia (K. Presi) litis & Corn<

4(1):126

subg. Breyniastrum (DC.) litis, comb. nov. — 4(1 ):1 24
calciphila (Standi. & Steyerm.) litis & Cornejo, comi
nov.— 4(1):126

cynophallophora f. linearifolia litis, f. nov— 4(1):98
domingensis (Spreng. ex CKI.) litis
nov.-4(l):129

domingensis subsp. grisebachii (Eichler) litis & Cornejo,
comb.nov— 4(1):129
dressleri Cornejo & litis, sp. nov. — 4(1):77
ferruginea (L.) litis & Cornejo, comb. nov. — 4(1 ):1 30
ferruginea subsp. cubensis (R. Rankin) litis & Cornejo,
comb.nov.-4(1):130

filipes (Donnell Smith) litis & Cornejo, comb, nov—
4(1):126

incana subsp. yucatanensis (Lundell) litis, comb, et stat.
nov— 4(1):83

indica (L) litis & Cornejo, comb, nov— 4(1):126
subg. Intutis (Raf.) litis, comb, et stat. nov— 4{1):128
isthmensis subsp. glabripetala Cornejo & 111
nov— 4(1):111

isthmensis subsp. mexicana Cornejo & lit

jamaicensis f. longifolia (Sw.) litis, comb, et stat. nov—
4(1):98

lindeniana Cornejo & litis, sp. nov— 4(1):83
lundellii (Standi.) litis & Cornejo, comb, nov— 4(1):1 27
mirifica (Standi.) litis & Cornejo, comb, nov— 4(1); 127
morenoi Cornejo & litis, sp. nov— 4(1):86
morenoi f. hastata litis, f. nov— 4(1);90

is & Cornejo, comb, nov— 4(1);1 27
D.nov— 4(1):104

a (L) litis & Cornejo, comb, nov— 4{1):98
singularis (R. Rankin) litis & Cornejo, comb, nov—
4(1):131

steyermarkii (Standi.) litis & Cornejo, comb, nov—
4(1):128

Salvia coriana Quedensley & Veliz, sp. nov — 4(1):27

V— nov— 4(1 ):20

Solanum

adenobasis M. Nee & Farruggia, sp. nov— 4(2):598
cyathophorum M. Nee & Farruggia, sp. nov — 4(2):596
Stenotis greenei (A. Gray) Terrell & H. Rob., comb, nov—
4(2):622

Stylidium darwinii Punekar & Lakshmin., sp. nov— 4(1):69

Tessiera hexasepala (Borhidi & Lozada) R.M. Salas & E.L
Cabral, comb, nov— 4(1 ):1 84

us var. pedunculatus (M.E. Jones)
ov.-4(l):207

epipsila var. repens (Turcz exTrautv. & CA. Mey.) R J. Little,
stat. nov— 4{1):225

pedatifidavar. brittoniana (Pollard) RJ. Little & LE. McKin-
ney, stat. nov.-4(l):225

praemorsa var. flavovirens (Pollard) R.J. Little, stat.
nov— 4(1);225

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SBM 12 I The “El Cielo” Biosphere Reserve, Tamaulipas, Mexico - $5.00
SBM 13 I Flora de Manantlan - $15.00

SBM 14 I Niebla & Vermilacinia (Ramalinaceae) from California & Baja California - $15.00
SBM 15 I Monograph of Northern Mexican Crataegus (Rosaceae, subfam. Maloideae) - $10.00
SBM 16 I Shinners & Mahler’s Illustrated Flora of North Central Texas - $89.95 (excluded from sale)
SBM 17 I The Grasses of Barbados (Poaceae) - $10.00

SBM 18 I Floristics in the New Millenium: Proceedings of the Flora of the Southeast U.S. Symposium - $5.00
SBM 19 I Emanuel D. Rudolph’s Studies in the History of North American Botany - $20.00
SBM 20 I Generic Conspectus of the Tribe Asterae (Asteraceae) in North & Central America - $8.00
SBM 21 I A Comparative Checklist of the Plant Diversity of the Iwokrama Forest, Guyana - $10.00
SBM 22 I Lloyd Herbert Shinners: By Himself - $14.00
SBM 23 I Taxonomy, Distribution, & Ecology of the Genus Phaseolus (Leguminosae) - $20.00
SBM 24 I Atlas of the Vascular Plants of Texas, Volume 1 & 2 - $25.00 (Vol. 1) I $15.00 (Vol. 2) | $30.00 (Set)
SBM 25 I Los Generos de Leguminosas del Norte de Mexico - $12.50
SBM 26 I Illustrated Flora of East Texas - $89.95 (excluded from sale)

SBM 27 I The Genus Psychotria (Rubiaceae) in the Philippine Archipelago - $18.00
SBM 28 I Wild Flowers of Mombacho (Nicaragua) - $10.00
SBM 29 I Muhlenbergia (Poaceae) de Chihuahua, Mexico - $12.50
SBM 30 I Tundra to Tropics: The Floristic Plant Geography of North America - $10.00
SBM 31 I A Fifth Checklist of Tennessee Vascular Plants - $15.00
SBM 32 I Gramineas de Zacatecas, Mdxico - $20.00

MISCELLANEOUS PUBLICATIONS

Keys to the Vascular Plants of the Black Gap Wildlife Management Area - $2.00
Mosses of Texas: A Manual of the Moss Flora - $7.50
Eriocaulaceae of Continental North America, North of Mexico - $2.00
Trees in the Life of the Maya World - $30.00
Violets (Viola) of Central & Eastern United States: An Introductory Survey - $2.00
The Ecology of our Landscape: The Botany of Where we Live - $7.50